A NEW SPECIES OF CAREBARA FROM THE PHILIPPINES WITH NOTES AND COMMENTS ON THE SYSTEMATICS OF THE CAREBARA GENUS GROUP (HYMENOPTERA: FORMICIDAE: MYRMICINAE)

2010·6.5 MB·English

by  Fernandez, Fernando

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A NEW SPECIES OF CAREBARA FROM THE
PHILIPPINES WITH NOTES AND COMMENTS ON THE
SYSTEMATICS OF THE CAREBARA GENUS GROUP
(HYMENOPTERA: FORMICIDAE: MYRMICINAE)
Una especie nueva de Carebara de Filipinas con notas
y comentarios sobre la sistemática del grupo Carebara
(Hymenoptera: Formicidae: Myrmicinae)
FERNANDO FERNÁNDEZ
Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Apartado 7495, Bogotá
D. C., Colombia. [email protected]
ABSTRACT
A new myrmicine ant species from the Philippines, Carebara alperti n. sp., is
described with taxonomic notes about the genus. The new species is congruent with
the genus defi nition proposed recently by Fernández (2004). The myrmicine ant genus
Parvimyrma is synonymized under Carebara (n. syn.), and the new combination
Carebara sangi (((EEEggguuuccchhhiii &&& BBBuuuiii 222000000777))) iiisss ppprrrooopppooossseeeddd... RRReeeiiinnnttteeerrrppprrreeetttaaatttiiiooonnn ooofff CCaarreebbaarraa
villiersi, originally described in the genus Nimbamyrma, suggests that Pheidologeton
should probably be synonymized within Carebara. A synoptic sketch of the systematic
of the CCCaaarrreeebbbaaarrraaa ggeennuuss ggrroouupp iiss pprroovviiddeedd ttoo sshhooww pprroobbaabbllee pphhyyllooggeenneettiiccss rreellaattiioonnsshhiippss..
Carebara arabara is proposed as a new name for Carebara striata Fernández,
2004:228.
Key words. Ants, Parvimyrma, synonymy, taxonomy
RESUMEN
Se describe una hormiga nueva de la subfamilia Myrmicinae de Filipinas, Carebara
alperti, n. sp., y se ofrecen notas taxonómicas sobre el género. La especie nueva es
congruente con la delimitación propuesta por Fernández (2004). El género Parvimyrma
es puesto en sinonimia con Carebara, proponiéndose la nueva combinación Carebara
sangi (Eguchi & Bui 2007). Se reinterpreta Carebara villiersi, descrita originalmente
como Nimbamyrma, con la sugerencia de poner en sinonimia Pheidologeton con Ca-
rebara. Se ofrece una sinopsis de la sistemática de las hormigas del grupo de géneros
Carebara junto con sus posibles relaciones fi logenéticas. CCCaaarrreeebbbaaarrraaa aaarrraaabbbaaarrraaa ssee pprrooppoonnee
como nuevo nombre para Carebara striata Fernández, 2004:228.
Palabras clave. Hormigas, Parvimyrma, sinonimia, taxonomía
INTRODUCTION (2003) proposed the solenopsidine tribe
group to cover the tribes Solenopsidini and
Among the myrmicine ants in need of Stenammini, and divided the Solenopsidini
phylogenetic studies are those grouped tribe into the Solenopsis and Carebara
around the Solenopsis Westwood and genus groups. The former corresponds
Carebara Westwood genus group. Bolton to the Solenopsidini tribe sensu Bolton
191
A new species of Carebara from the Philippines
(1987), and the latter to the Pheidologetini proposal of Fernández (2004:196), Carebara
tribe in its original sense (e.g. Ettershank appears broader and probably monophyletic.
1966). Currently, there are no published In this publication a new species of Carebara
phylogenetic studies that can corroborate from the Philippines is described. This
or reject this proposal, although ongoing species has mixed attributes that recall ants
studies suggest that the Solenopsidini tribe of the Carebara escherichi and Carebara
is not a natural group (Rodríguez et al., in concinna groups, and is consistent with
preparation) and further suggest a tendency the Carebara generic concept proposed in
towards parallel evolution in some traits Fernández (2004).
in these ants, including clypeal reduction
between antenal lobes, reduction of the eyes Additional notes about Parvimyrma Eguchi
to entirely absent or with few ommatidia, & Bui and Pheidologeton Mayr are also
reduction in the number of antennal and provided. The recently described genus
palpal segments, and perhaps, simplifi cation Parvimyrma is hereto merged with Carebara.
in sculpture. Part of this can be due to Parvimyrma sangi is a Carebara species with
adaptation to similar habits (e.g. underground central clypeal hair, an attribute sporadically
habits) or to miniaturization (as in several present in other Carebara workers (see
Solenopsis species or in the Carebara lignata below).
group), perhaps associated in part with
lestobiotic (opportunistic thievery) habits. Adjustments to the description of Carebara
(= Nimbamyrma) villiersi are also given. The
Moreover, members of the Carebara genus latter species seems to be a bridge between
group exhibit complex biology that has not Carebara and Pheidologeton. The status of
been studied. Subjects requiring further Pheidologeton is weak, because this genus
investigation include the intercastes, the does not have attributes that clearly separate
disappearance of the intermediate caste, major it from Carebara (Fernández 2004). Further
workers of exaggerated size (and big queens) investigation is likely to show if that this genus
occurring with very small blind minor workers should be merged under CCCaaarrreeebbbaaarrraaa aanndd rreedduucceedd
(some less than 1mm in total length). The to a group of species within that genus.
genetic and physiological bases of some of
these attributes are not understood, as well as Finally, I present a sketch of the taxonomy and
the natural history of most of the species. hypothesis to test the phylogeny of the ants
in the Carebara genus group, as a working
The Carebara group includes dimorphic and synopsis to orient future investigations in the
secondarily monomorphic ants with some systematics of these ants.
species displaying an acute dimorphism
of size between the female and the worker MATERIALS AND METHODS
(Hölldobler & Wilson 1990). Around this
genus have been described others like Measurements were made using a Leica
Erebomyrma Wheeler or Oligomyrmex Mayr. stereomicroscope at 80X magnifi cation and
Although characters used for inclusion of a fi ber ring lamp. All measurements are in
species in these genera seemed consistent, mm:
the discovery of new species and the
reinterpretation of others force a broader HL Head Length. The length of the head
circumscription of Carebara, including capsule excluding the mandibles;
Oligomyrmex and other genera as junior measured in full-face view, as a straight
synonyms (Fernández 2004). In the generic line from the mid-point of the anterior
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Fernández
clypeal margin to the mid-point of the processing using Adobe Photoshop CS.
posterior margin. Figures 1C and 1F were taken from the “Ants
HW Head Width. The maximum width of the of Philippines” web page hosted by Gary
head behind the eyes, measured in full- Alpert, David General and Ven Samarita
face view. (www.discoverlife.org/mp). Figures 2A, B
EL Eye Lenght (queen and male). Maximun were taken from Fernández 2004:207.
diameter of compound eye.
ML Mandible Length (queen and male). Specimens studied and deposited in the
In full face view, the maximum length following collections:
between anterior clypeal margin and
mandible apex, with mandibles closed. ICN. Insect Collection, Instituto de Ciencias
SL Scape Length. The maximum length of the Naturales, Universidad Nacional de Colombia,
scape, excluding the basal constriction or Bogotá D.C., Colombia.
neck that occurs just distal of the condyle
bulb. MCZC. Museum of Comparative Zoology,
PW Pronotal Width. The maximum width Harvard University, Cambridge, USA.
of the pronotum in dorsal view. Worker
only. SYSTEMATIC TREATMENT
WL Weber’s Length of Mesosoma. The
diagonal length of the mesosoma in Carebara alperti, new species
profi le, from the anteriormost point of the Figures 1A-1G
pronotum to the posterior basal angle of
the metapleuron. Worker measurements (holotype): HL 0.52
PL Petiole length, in lateral view. HW 0.48 SL 0.40 PW 0.30 WL 0.54 PL 0.19
PPL Postpetiole length, in lateral view. PPL 0.10 GL 0.65 TL 2.04 CI 92 SI 83.
GL Gaster length, in lateral view.
TL Total Length. The total outstretched Head slightly longer than wide. Posterior
length of the ant from the mandibular cephalic border sinuous, lateral sides slightly
apex to the gastral apex, that is, convex. Mandibles conspicuous with four
HL+ML+WL+PL+PPL+GL. stout teeth. Median portion of clypeus
CI Cephalic Index. (HW/HL)*100 bicarinate. Eyes with one ommatidium,
SI Scape Index. (SL/HW)* 100. situated anterior to cephalic midline. Antennae
11-segmented with a 2-segmented apical club.
Ants were mounted on black, archival quality Scapes failing to reach the vertexal border in
paper points. Pictures of the ants were taken less than their maximum width.
for the author in the Ant Room, MCZC.
The imaging system consisted of Leica Promesonotum, in profi le, strongly convex.
MZ16 stereo microscope, motor focus drive Propodeum convex and low, unarmed.
and JVC KY-F70B digital camera. Basic Propodeal spiracle relatively small, circular,
Apochromatic magnifi cation range is 7.1x high and equidistant from propodeal border.
to 115x with a 10x eye piece correction Propodeal lobes small. Petiole with long
for accurate color rendition. Lighting was peduncle and with a well-defi ned high node;
via three fluorescent lights and a velum petiolar spiracle half of petiolar length.
paper light box to create even lighting. Dell Subpetiolar process absent. Postpetiole
Windows computer with Auto-Montage dorsally convex, lower than petiole.
Professional by Syncroscopy Final image Postpetiole, in dorsal view, trapezoidal.
193
A new species of Carebara from the Philippines
A B
C D
E F
G
Figure 1. Carebara alperti new species. A. Holotype worker, head in frontal view. B. Holotype
worker, lateral view. C. Queen paratype, head in full face view. D. Queen paratype in lateral
view. E. Male paratype in lateral view. F. Male paratype, head in full face view. G. Male
genitalia. Figures 1C and 1F from Ants of Philippines Web Page.
194
Fernández
Sting well developed. Body smooth and hairs on rest of head. Body dark brown,
shiny, except for mesopleura and sides of appendages, antennae, and mandibles
propodeum, which are foveated. Dorsum of brown.
petiole and postpetiole smooth and shining.
Erect hairs absent, except for some few hairs Holotype worker: Philippines, Negros
in the last gastral tergum. Short apressed hairs Oriental, Dumaguete, Horns of Negros,
sparse on body. Body dark brown, appendages Camp Lookout, 10 ix 1948, J. W. Chapman
lighter brown. leg. No. JWC0002 (deposited in MCZC).
Paratypes (same data): one worker (deposited
Female measurements. HL 1.35 HW 1.40 EL in ICN), one queen and one male (deposited
0.38 ML 0.25 SL 0.75 WL 2.48 PL 0.73 PPL in MCZC).
0.55 GL 3.42 TL 8.78 CI 103 SI 54.
Comments. This species can be differentiated
As typical myrmicine queen, although from other CCCaaarrreeebbbaaarrraaa bbyy tthhee ccoommbbiinnaattiioonn ooff tthhee
noticeable larger than worker (Figures 1C-D). following traits: eyes present, promesonotum
Head wider posteriorly, widest point near to strongly convex, propodeum unarmed and
occipital corner. Vertexal border with median propodeal lobes small. The presence of eyes
concavity. Clypeal border evenly convex. links this species with the Carebara concinna
Antenna segmented. Scapes short, widening group and the propodeum with the Carebara
distally. Ocelli well defi ned. Propodeum with escherichi group. The promesonotum clearly
two well-defi ned, strong spiniform process. convex appears as unique in Carebara. This
Most of body strongly smooth and shining. species is placed, provisionally, in his own
Head with conspicuous longitudinal rugulae, species group (Fig. 5).
except the central clypeal area. Front wing
with marginal, fi rst submarginal and fi rst This species is dedicated in honor to the
discoidal cells present and closed. Vein M colleague Gary Alpert (MCZC), for their
ending near to wing margin.Very few erect generosity and helping to the visiting
hairs on head frons, and dorsal masticatory myrmecologists in Cambridge, and for their
border of mandibles, sparsed erect hairs on kindly advice with the specimens collected
mesosoma, petiole, postpetiole, and gaster and AutoMontage pictures.
(especially fi rst tergum). Body dark brown;
appendages, antennae, and mandibles TAXONOMIC NOTES IN CAREBARA
brown.
Parvimyrma Eguchi & Bui, 2007, is
Male measurements. HL 0.80 HW 0.98 EL junior synonym of Carebara (n. syn.),
0.38 ML 0.15 SL 0.18 WL 1.88 PL 0.60 PPL corresponding to the species Carebara sangi
0.38 GL 2.38 TL 6.19 CI 123 SI 18. (n. comb.), from Vietnam. As pointed out
by authors (Eguchi & Bui, 2007:42) “... the
As typical myrmicine male (Figures 1E- presence of mediate clypeal setae is the only
G). Mandibles with 5 well-defi ned teeth, characteristic separating Parvimyrma from
decreasing in size from apex. Promesonotum Carebara” (italics mine). In fact, C. sangi
convex in lateral view. Propodeum angulated is a typical Carebara of the lignata group
in lateral view. Gonostylus pale, with dense whose workers have a central hair as an
pilosty of curved withish hairs; volsellae apomorphic trait. It is true that the central
dark, elongated, ending in a rounded apices. hair is absent in the basic plan of Carebara,
Abundant white erect hairs on mandibles, but the author has discovered several workers
clypeus, and gena, dense black erect short of C. lignata (two examples in Figs. 2D
195
A new species of Carebara from the Philippines
& E) with central clypeal hair, in addition The Carebara genus group, as proposed by
to the case of C. peruviana (Fernández Bolton (2003) includes the genera Adlerzia
2004). Furthermore, the study of abundant Forel, Carebara Westwood, Machomyrma
material of Carebara in MCZC shows that Forel, Mayriella Forel, Pheidologeton Mayr
the clypeal hairs can be variously displayed. and Tranopelta Mayr.
We still do not know the genetic basis of this
variation, but the observations suggest that The Carebara genus group (except Mayriella,
is very inappropiated to create a new genus see below) can be defi ned by the following
based only by a feeble trait. Parvimyrma traits (based partially on Bolton 2003):
does not have any other characteristic or clypeus constricted posteriorly, narrowly
set of attributes that allow it to be separated inserted between frontal lobes; antennae 9
clearly from Carebara. The authors place to 11 (rarely 8) -segmented with antennal
this genus in the Solenopsis group, but, club 2 or 3- segmented; antennal sockets
except the possession of the central clypeal and inner margins of frontal lobes in close
hair, there are not strong arguments for such proximity; median portion of clypeus
placement. usually bicarinate; clypeus always with a
pair of setae, commonly a second pair of
The internal taxonomy and phylogeny of setae (“paracarinal” in Eguchi & Bui 2007
Carebara have been partially explored terminology); clypeus usually without an
(taxonomy) or remains practically unknown isolated clypeal seta (sporadically present);
(phylogeny). The small size is one of the main major workers, when present, with long
factors that have discouraged researchers. heads; polymorphic to monomorphic.
To this we must add monotony in external
traits in the smaller workers and, overall, This proposal does not include Mayriella,
the existence of isolated samples of workers a specialized ant that is probably not a
without majors, females and associated member of the Carebara genus group. In
males (e.g nidotypes). The taxonomy of this genus the antennae is 10-segmented
the American species has been studied with a 2-segmented club, palpal formula
(Fernández 2004) and the Malagasy and 4,3; the clypeus is concave in the middle
Ethiopian fauna is underway. Nevertheless, and laterally bidentate; and the entire body
studies of the Asiatic and Australian faunas is coarsely sculptured. In the carebarine ants
are needed. In total there are some 180 the antennae+club combination 10 + 2 is
species described in the world, the majority very rare, and no other ant in this group has
in the Southern Hemisphere. a palpal formula of 4,3 segments associated
with this antennae confi guration. Also, no
PHYLOGENY AND SYSTEMATICS IN carebarine has the clypeus so modifi ed as
CAREBARA GENUS GROUP in Mayriella, nor the coarse sculpture of the
body, especially in the head.
The proposed classification of the
solenopsidine ants of Bolton (2003) Some traits in the Carebara genus group are
has not been evaluated in phylogenetic discussed below.
studies. The tribe Solenopsidini seems to
be non-monophyletic (Rodriguez et al., in Antennae. All members of the group have 8
preparation) based on morphological grounds. to 11 segments on antennae (a reduction from
We need morphological and molecular the basic number of 12, which is probably
studies that evaluate the phylogenetic status plesiomorphic in Formicidae, see Bolton
of the Solenopsis and Carebara genus group. 2003:288). This reduction has occurred several
196
Fernández
times in the Myrmicinae. On the one hand, upward and outward. In many workers of
all the carebarines have 2- or 3- segmented the Carebara group a second pair of hairs
antennal clubs. It is diffi cult to determine if a exists (called paracarinal setae in Eguchi &
2- or a 3- segmented club is plesiomorphic. Buy 2007). In many workers of Carebara
It is assumed that a multisegmented club the central pair and paracarinal hairs are
is plesiomorphic by its presence in several distinguished clearly from other clypeal hairs.
lineages of Hymenoptera Apocrita. On the As has been documented for other traits, these
other hand, ants with specialized habits and hairs attenuate or disappear in major workers
morphology, like Discothyrea Roger, present and queens. For now, we lack comparative
a single segmented club. In Solenopsidini, studies of the ontogeny and morphology
Solenopsis, an apparently derived genus, has of the median setae in ants. The median
a 2-segmented club. setae appears to be similar in position in
solenopsidines, whereas in different position
Palps. From the plesiomorphic number of in attines (Brandão & Mayhe-Nunes 2001).
6,4 palps in Hymenoptera and other groups,
reduction in the number of palpal segments Caste. The carebarines include polymorphic,
have occurred several times in Formicidae, dimorphic, and monomorphic ants.
even to zero in some cases. Adlerzia show Apparently the ancestral carebarine was di-
the most plesiomorphic number with or polymorphic, with an evolutionary trend
palpal formula 4,3 for the Carebara-group. of disappareance of the intermediate worker
Tranopelta displays a reduction to 3,2 and caste (Hölldobler & Wilson 1990) with tiny
the rest of the carebarines present a uniform smaller workers (in contrast to very large
reduction to 2,2. queens) in the species groups concinna,
crigensis, lignata and escherichi.
Bicarinate clypeus. This it is an attribute
common in the tribes Solenopsidini, Eyes. From the most basal to the most derived
Adelomyrmecini, and Stenammini (Bolton groups there is a tendency towards reduction
2003). The great majority of the ants on these on the number of ommatidia in the eyes from
tribes have a median bicarinate clypeus. In few in C. escherichi and C. crigeri species
Adelomyrmecini both carinae can merge groups or completely absent in C. lignata
forming a fused keel, as can be seen in species group.
Cryptomyrmex Fernández or Baracidris
Bolton. This trait is present in all minor Elongated heads in major workers. With the
workers, and is faded or secondarily absent in exception of Pheidologeton and Tranopelta,
major workers (in cases on which this caste all Carebara major workers have elongated
is present) and workers of several species of heads, with CI greater than 130 (Figs. 3A,B).
Pheidologeton. This trait is also present in other Myrmicinae
including some Pheidole (especially in the
Clypeal hairs. In the workers of the Carebara aberrans groups) and one undescribed species
genus group there is always a pair of setae in of Solenopsis from Argentina. Because these
the central part of the clypeal anterior margin latter taxa appear to be only distantly related to
(Fig. 2 A,B,C). Occasionally there are workers Carebara, the presence of an elongated head
with a central hair (C. anophtalma, C. lignata is here interpreted as a convergent trait. The
Fig. 2 D,E, P. sangi), but this must taken as presence of major workers with heads not so
an apomorphic attribute of these species. elongated in Pheidologeton (Fig. 3C), implies
This pair of hairs are always conspicuous the retention of a plesiomorphic trait or the
(rarely displaced from the center) and directed new acquisition of wide heads.
197
A new species of Carebara from the Philippines
A B
C D
E
Figure 2. Clypeal setae in Carebara workers. A & B: Carebara reticulata. A. Head in full
face view. B. Head in oblique frontal view. C. Carebara urichi. D & E. Central clypeal setae
in some Carebara lignata. A & B from Fernández (2004:207).
198
Fernández
A B
C
Figure 3. Head in major workers in some Carebara genus group. A. Adlerzia froggati. B.
Machomyrma dispar. C. Pheidologetonsilenus.
OUTLINE OF EVOLUTION IN THE if a true member of the Carebara group,
GROUP would represent a separate lineage, with
the loss of the major worker and palpal
An outline of evolution in the group reduction to 3,2. Machomyrma + Carebara
is sketched, with the intention of start s.l. present reduced palpal formula of 2,2. At
morphological and molecular phylogeny this node two lineages can be differentiated,
studies that can either confirm, reject or Machomyrma and Carebara sensu lato.
change the scheme proposed here. What Machomyrma (monotypic genus from
follows is a “working hypothesis” (Fig. Australia) would retain the plesiomorphic
5). The carebarine had 11- segmented club of 3 segments. Carebara sensu lato
antennae with 3- segmented club as probable (including Pheidologeton) is characterized
plesiomorphic state; palpal formula 4,3; by a 2-segmented antennal club.
and dimorphic castes, the major workers
with elongated heads. Adlerzia, the most In Carebara sensu lato some groups retain
probable plesiomorphic genus, appears as 11-segmented antennae (Pheidologeton),
the sister group of Machomyrma + Carebara 11-9 (C. concinna group) and other present
s.l. This is a monotypic taxon from Australia reduction to 10 segments (C. crigensis group),
that retains an antennal club of 3 segments 9-8 (C. escherichi group) and 9 (C. lignata
and palpal formula 4,3. Tranopelta (with group); whereas Pheidologeton retains the
two species confined to the Neotropics), major worker with wide heads (or this trait
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A new species of Carebara from the Philippines
maybe a novelty), the C. concinna group stings by Kugler (1986) suggests that some
(probably paraphyletic) retains dimorphic Pheidologeton are more closely related to
castes (both major and minor workers with Oligomyrmex than to other Pheidologeton.
eyes) and the C. lignata group always with
eyeless minor workers, a unique trait in the THE FATE OF PHEIDOLOGETON
Carebara group. The other Carebara groups
are united by the disappearance of the major The wide heads of the major workers and
worker: C. crigensis group (monotypic) with caste polymorphism puts Pheidologeton in
mandibles with 2 teeth and C. escherichi a more diffi cult position in this scheme. This
group (formerly Paedalgus) with narrow genus appears to be closely related to, but
head and short propodeum. differentiated from, Carebara. However,
C. villiersi (see below) could suggests the
The ants of the lignata group deserve placement of Pheidologeton as member of
separate mention. Members of this species the Carebara sensu lato (Fig. 5).
group are in agreement with the concept of
Carebara s. str. of earlier literature (Bolton The antennal and palpal formula puts
2003), that is, all minor workers lacking Pheidologeton within Carebara, although
eyes. In Fernández (2004) it is clear that separated from Carebara by the presence
some of these Carebara have, in addition of polymorphism in all his species. Also,
to blindness, small workers, eyed major the pair of clypeal setae in the smaller
workers (or at least with one ommatidium). workers is absent in several species. The
Due to the difficulty in collecting these polymorphism can be interpreted as an
subterranean ants, there is a small quantity independent acquisition in this lineage,
of major workers in museum collections. from a dimorphic ancestor (ancestor of
On the other hand, it is in this group that Machomyrma + Carebara), as well as the
the smallest ants in the world are known, undifferentiated clypeal pair of setae in some
with some species barely approaching one species.
millimeter in total length or 0.21 mm in
head width. Nimbamyrma villiersi, described from
workers of Guinea by Bernard (1953), plays an
As pointed out by Fernández (2004), the important role in the taxonomy of Carebara.
synonymy of the genera Oligomyrmex and The lack of observed material of this species
Paedalgus with Carebara seems inadmissible prevented a defi nitive evaluation of the genus
at fi rst view. Nevertheless, the existence of in Fernández (2004). Nevertheless, recent
several “bridge” species obscures the limits material sent to the author by Barry Bolton,
between these genera and Carebara. Several and observation of workers at MCZC has
species have major workers that belong allowed better study of the propodeal teeth
to the generic concept of Oligomyrmex and comparison to Bernard’s description.
whereas their smaller workers are Carebara According to this author, the propodeum, in
s.str. Carebara intermedia, from Trinidad, lateral view, has two kinds of strong teeth:
has attributes of both the C. escherichi the propodeal teeth and the “inferior ones”,
and C. concinna groups. Carebara alperti which correspond to the metapleural process.
(described above) has a mixture of traits Actually, the metapleural process forms an
of the C. escherichi group and C. concinna angulated process, but less conspicuous than
group. Carebara villiersi (see below) was described and illustrated in Bernard
resembles Pheidologeton. The study of ant (1953).
200