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A New Social Parasite in the Ant Genus Ectatomma F. Smith (Hymenoptera, Formicidae, Ectatomminae)

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Ectatomma parasiticum Feitosa & Fresneau, a new ant species socially parasitic on Ectatomma tuberculatum (Olivier), is described from gynes discovered in Apazapan, state of Veracruz, Mexico, and reared in the laboratory. Ectatomma parasiticum is the first social parasite described in the Ectatomminae. This species can be distinguished from its host by morphological and behavioral features characteristic of the inquilines known in other ant subfamilies including reduced size, thickened petiole, and agonistic interactions with host species.
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Accepted by J.Longino: 17 Jan. 2008; published: 27 Feb. 2008 47
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2008 · Magnolia Press
Zootaxa 1713: 4752 (2008)
www.mapress.com/zootaxa/
A new social parasite in the ant genus Ectatomma F. Smith
(Hymenoptera, Formicidae, Ectatomminae)
RODRIGO M. FEITOSA1, RIVIANE R. HORA2, 3, JACQUES H. C. DELABIE4, 5, JORGE VALENZUELA6
& DOMINIQUE FRESNEAU2
1Museu de Zoologia da Universidade de São Paulo, Av. Nazaré 481, 04263-000, Ipiranga, São Paulo, SP, Brazil.
E-mail: rfeitosa@usp.br
2Laboratoire d’Ethologie Expérimentale et Comparée (CNRS UMR 7153), Université Paris-Nord, 99, Avenue J. -B. Clément, 93430,
Villetaneuse, France. E-mail: Dominique.Fresneau@leec.univ-paris13.fr
3Departamento de Biologia Geral, Universidade Federal de Viçosa, Av. P.H. Rolfs s/n, 36570-000, Viçosa, MG, Brazil.
E-mail: rivianer@hotmail.com
4Laboratório de Mirmecologia, Convênio UESC/CEPEC, Centro de Pesquisas do Cacau, C.P. 7, 45600-000, Itabuna, BA, Brazil.
E-mail: delabie@cepec.gov.br
5Departamento de Ciências Agrárias e Ambientais, Universidade Estadual de Santa Cruz, 45650-000, Ilhéus, BA, Brazil.
6Instituto de Ecologia, A. C., Km 2.5, antigua carretera a Coatepec, A.P. 63, 91000, Xalapa, Veracruz, Mexico.
E-mail: jorge.valenzuela@inecol.edu.mx
Abstract
Ectatomma parasiticum Feitosa & Fresneau, a new ant species socially parasitic on Ectatomma tuberculatum (Olivier),
is described from gynes discovered in Apazapan, state of Veracruz, Mexico, and reared in the laboratory. Ectatomma
parasiticum is the first social parasite described in the Ectatomminae. This species can be distinguished from its host by
morphological and behavioral features characteristic of the inquilines known in other ant subfamilies including reduced
size, thickened petiole, and agonistic interactions with host species.
Key words: Ectatomminae, Ectatomma tuberculatum, Social parasitism, Taxonomy, Mexico
Introduction
The ant genus Ectatomma includes 14 relatively large species occurring predominantly in the Neotropical
Region, with the widespread species Ectatomma tuberculatum (Olivier) also occurring in the Neartic Region
(Bolton et al. 2006; Fernández & Ospina 2003). These ants occur in a variety of warm habitats, and because
they are often abundant and conspicuous, they figure prominently in ecological studies (Kugler & Brown
1982). Species of Ectatomma are generalized predators of a variety of small arthropods and earthworms in
addition to collecting honeydew from homopterous insects and nectar from plant sources (e.g. Weber 1946;
Wheeler 1986; Dejean & Lachaud 1992; Pie 2004).
Instances of social parasitism in ants have a patchy taxonomic and geographic occurrence, being well
known among north temperate ants in the subfamilies Formicinae and Myrmicinae (Wilson 1971; Hölldobler
& Wilson 1990). Hora et al. (2005) demonstrated the parasitic nature of microgynes found in colonies of a
Mexican population of E. tuberculatum. These individuals concentrate their reproductive efforts almost exclu-
sively on the production of sexual offspring, and they are genetically distinct and reproductively isolated from
their hosts.
FEITOSA ET AL.
48 · Zootaxa 1713 © 2008 Magnolia Press
In the present paper we describe the microgyne from Hora et al. (2005) as a new species, Ectatomma par-
asiticum Feitosa & Fresneau, a social parasite. It is the fifteenth species described in the genus and represents
the first case of social parasitism in the subfamily Ectatomminae.
Material and methods
The species described in the present study was obtained while collecting colonies of E. tuberculatum during
fieldwork in Apazapan, in the state of Veracruz, Mexico (19º19’38”N; 96º43’21”W) from September 1999 to
July 2000. The parasitic gynes were promptly distinguished by eye on the basis of their reduced size com-
pared to E. tuberculatum gynes. Colonies were reared in laboratory and new parasitic specimens were pro-
duced (about 65 gynes from eight different colonies). Depository collections are referred to by the following
acronyms:
CPDC Centro de Pesquisas do Cacau, Itabuna, Bahia, Brazil.
INEC Instituto de Ecologia, Xalapa, Veracruz, Mexico.
LACM Los Angeles County Museum of Natural History, Los Angeles, California, USA.
MZSP Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil.
UNCB Museo de Historia Natural del Instituto de Ciencias Naturales, Bogotá, Colombia.
USNM National Museum of Natural History; Smithsonian Institution, Washington, DC, USA.
Scanning Electron Microscope (SEM) images of Ectatomma parasiticum were made at MZSP with a
LEO 440® microscope. The specimen was previously cleaned in acetone, critical-point dried in a Balzer (Bal-
Tec® CPD 030), and coated with gold (Bal-Tec® SCD 050). Measurements were obtained with a micrometric
reticule and using the scale of the SEM. All measurements are given in mm and the abbreviations used are:
HL head length; the maximum measurable length of head capsule excluding mandibles, measured in
full-face view, in a straight line from the mid-point of the anterior clypeal margin to the mid-point of
the vertexal margin.
HW head width; the maximum width of the head capsule measured in full-face view, excluding the com-
pound eyes.
SL antennal scape length; the chord length of the antennal scape, excluding the basal condyle and its
peduncle.
EL eye length; the maximum measurable length of eyes in profile.
PW pronotal width; in dorsal view, the maximum width of the pronotum.
WL mesosoma length (Weber´s length); the diagonal length of mesosoma in profile, from the mid-point
of the anterior pronotal declivity to the posterior basal angle of the metapleuron.
PTL petiole length; in dorsal view, the maximum length of the petiole.
PTW petiole width; in dorsal view, the maximum width of the petiole.
CI cephalic index. HW x 100/HL.
SI scape index. SL x 100/HW.
OI optical index. EL x 100/HW.
PTI petiolar index. PTL x 100/PTW.
Zootaxa 1713 © 2008 Magnolia Press · 49
A NEW SOCIAL PARASITE IN ECTATOMMA
Results
Ectatomma parasiticum Feitosa & Fresneau, sp. nov.
Figures 1, 2
Holotype gyne. MEXICO: Apazapan, Veracruz, 19º19’38”N 96º43’21”W, ix.1999, D. Fresneau col. [INEC].
Paratypes. same data as holotype (1 gyne) [CPDC]; (1 gyne) [UNCB]; (1 gyne) [INEC]; vii.2000, D.
Fresneau & R. Hora cols. (1 gyne) [LACM]; (2 gynes) [MZSP]; (1 gyne) [USNM].
Diagnosis. Size relatively small (WL approximately 3.80 mm); clypeus and frontal area without sculp-
ture; antennal scapes longer than the maximum head width (SI > 108); petiole relatively thick in lateral view.
Gyne description. Holotype (paratypes): HL 2.10 (2.06–2.16); HW 1.85 (1.73–1.88); SL 2.04 (1.92–
2.06); EL 0.51 (0.50–0.58); PW 1.69 (1.62–1.77); WL 3.88 (3.65–4.04); PTL 0.79 (0.78–0.88); PTW 0.92
(0.92–1.07); CI 88.07 (84.11–88.89); SI 110.42 (108.57–113.04); OI 27.50 (27.50–31.25); PTI 85 (82.14–
85.71). Color yellowish brown to dark reddish brown, including appendages. Mandibles finely and densely
striate, with sparse piligerous punctures; clypeus, genae, and frontal area predominantly smooth, but opaque;
dorsal surface of head densely and coarsely reticulated, except for the areas of antennal articulations, which
are finely punctate; ventral surface of head with sparse longitudinal striae; antennal scapes finely and longitu-
dinally striate. Mesosoma with variously oriented costulae, from sparse and transverse on dorsum of prono-
tum and propodeum to dense and subconcentric on the dorsum of scutum and scutellum; forecoxae with
dense, fine, regular transverse striation; legs mostly smooth and shining. Lateral and posterior faces of petiolar
node with sparse, short, longitudinal costulae; sculpture of gaster consisting of arched, transverse costulae,
becoming gradually finer from first to terminal segment.
Pilosity cream-colored. Body covered by relatively sparse, long, suberect hairs; antennal scapes and legs
with short, suberect hairs; antennal funiculi and tarsi covered by fine apressed pubescence.
Head subrectangular, with weakly convex lateral borders and vertexal margin straight; masticatory mar-
gins of mandibles multidenticulate and with a large apical tooth; clypeus strongly convex anteriorly; frontal
lobes reduced; scapes in repose fairly surpassing the posterolateral margins of vertex; funicular segments
gradually thickened distally; compound eyes placed near the posterolateral portions of head; ocelli present
and reduced in size.
Pronotum with a distinct median eminence directed forward and a conspicuous pair of dorsolateral
(humeral) projections; scutum large and rounded; notauli almost indistinct among sculpturation; parapsidial
lines feebly visible and subparallel; scutoscutellar sulcus deeply impressed; scutellum relatively narrow and
strongly convex, in lateral view; dorsal face of propodeum meeting the declivous face in a pair of reduced,
blunt teeth; propodeal spiracle elliptical. Wing venation fully developed. Forewing with a weakly colored
stigma; longitudinal veins Sc+R, SR, M, Cu, and A present; SR extending distally beyond stigma, forming 1R
and 2R cells; cross vein 1r vestigial, not forming the 2R cell; M and Cu also extend distally as tubular veins
for most of their length; A not extending beyond the junction with Cu; C, R, Cu, 1M, 1Cu, and SR cells
closed. Hind wing with Sc+R extending beyond point where they connect to M, which continues as a tubular
vein as much as Sc+R and then extends as spectral vein to wing distal border; basally M+Cu extending as a
tubular vein beyond junction with Anal vein, which continues shortly beyond this point; seven submedian
hamuli present.
Petiole ventrally carinate; in lateral view, petiolar node thick and subtriangular; anterior slope nearly con-
cave and posterior slope slightly convex. Sternite of first gastral segment with a distinct anterior projection.
Worker. Unknown (but see comments bellow).
Male. Unknown.
Etymology. The specific epithet is a reference to the parasitic nature of this species.
FEITOSA ET AL.
50 · Zootaxa 1713 © 2008 Magnolia Press
FIGURE 1. Paratype gyne of Ectatomma parasiticum: A, head in full face view; B, lateral view; C, dorsal view.
Comments. Gynes of the socially parasitic Ectatomma parasiticum can be distinguished from the gynes
of its host species, E. tuberculatum, by the following features: sparser sculpture on the body; smaller size (Fig.
2), with WL approximately 3.80 mm (around 5.40 mm in E. tuberculatum); clypeus and frontal area devoid of
any sculpture (usually longitudinally striate in E. tuberculatum); antennal scapes longer than the maximum
head width, with SI > 108 (< 99 in E. tuberculatum); propodeal spines reduced to minute teeth; and petiole
Zootaxa 1713 © 2008 Magnolia Press · 51
A NEW SOCIAL PARASITE IN ECTATOMMA
thicker in lateral view (flattened anteroposteriorly in E. tuberculatum). The reduced size and widener petiole
of E. parasiticum are also characteristic of the inquiline syndrome in other ant species (Wilson 1984; Rad-
chenko & Elmes 2003).
Males produced by parasitized colonies were of a uniform morphology and indistinguishable from males
of E. tuberculatum. Thus it remains unknown if males of E. parasiticum are lacking, present but not yet
observed, or observed but indistinguishable from E. tuberculatum. According to Hora et al. (2005), one of the
10 colonies of E. parasiticum reared in laboratory produced four small "workers." However, these workers
presented a developed spermatheca and six to 10 ovarioles, in contrast to typical workers of E. tuberculatum
which lack the spermatheca and possess only one to four ovarioles (Fénerón & Billen 1996; Hora et al. 2001).
The presence of developed reproductive structures in these specimens suggests that they are possibly interme-
diate (intercaste?) reproductive forms of E. parasiticum and not true workers.
Up to now, the occurrence of this species is restricted to Apazapan, state of Veracruz, Mexico. However,
its host, E. tuberculatum, is widely distributed in the Neotropics, from Mexico to northern Argentina. We
expect that the excavation and detailed examination of E. tuberculatum colonies in different localities could
reveal new populations of E. parasiticum.
A detailed behavioral and genetic study on the interaction between E. tuberculatum and E. parasiticum
(so far undescribed and treated as “microgynes”) was conducted by Hora et al. (2005). Gynes and workers of
E. tuberculatum and gynes of E. parasiticum were sequenced for the cyt b region and the results showed two
haplotypes. The haplotypes differed in seven variable sites, with a nucleotide sequence difference of 0.93%.
They clearly discriminate E. parasiticum from the group composed of workers and gynes of E. tuberculatum.
According to the findings of Hora et al. (op. cit.), E. parasiticum is a genetically distinct social parasite pro-
ducing of almost exclusively sexual offspring. The co-occurrence of E. parasiticum and E. tuberculatum in
the field (nine mixed colonies found) suggests that the parasite usurps established colonies of the host, but
does not kill the resident gynes. Agonistic interactions were also observed, exclusively from workers and
gynes of E. tuberculatum against the parasites.
FIGURE 2. Morphometric scatterplot of head width by scape length, differentiating gynes of Ectatomma parasiticum
and E. tuberculatum. Measurements are in mm.
FEITOSA ET AL.
52 · Zootaxa 1713 © 2008 Magnolia Press
Microgynes are also found in Ectatomma ruidum; however, this is a truly gyne-polymorphic species, and
the offspring of both microgynes and normal gynes consist of workers, males, and both microgynes and nor-
mal gynes. It was therefore suggested that the two gyne morphs in E. ruidum represent alternative phenotypes
adapted to different ways of dispersal and colony founding. Ectatomma ruidum microgynes are thought to dis-
perse and to found new colonies solitarily, while the macrogynes are a stationary morph (Lachaud et al. 1999).
The study of Hora et al. (2005) and the present paper raise the possibility of the occurrence of similar
undescribed social parasites within other basal ant lineages.
Acknowledgements
The authors would like to thank Alex Wild, Alexander Radchenko, Riita Savolainen, and Jack Longino for the
critical reading and the invaluable comments on the manuscript. Lara M. Guimarães took the SEM images.
Rogério Rosa da Silva kindly prepared the morphometric scatterplot. We also acknowledge M. Favila, L.
Quiroz (INEC, Mexico), C. Rojas, A. Wolf, and K. Wolf for the technical support during fieldwork. RRH
received financial support from FAPEMIG and CAPES, Brazil, Project CAPES/COFECUB no. 244/98-II.
RMF and JHCD acknowledge the research grant received from FAPESP and CNPq, respectively. This work
was partially supported by BRI (Université Paris-Noird, France).
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... Some of these problems were partly solved by Kugler and Brown in 1982, who suggested some synonymies and recognized 12 species. Recently, three new species were described or redescribed (Almeida 1986(Almeida , 1987Arias-Penna 2006;Feitosa et al. 2008), but nothing is known on their relationships. ...
... Microgynes are present in species of the ant subfamilies Pseudomyrmecinae (Janzen 1973), Formicinae (Heinze and Hölldobler 1993), Myrmicinae (Elmes 1991;McInnes and Tschinkel 1995;Hamaguchi and Kinomura 1996;Schlick-Steiner et al. 2005;Lenoir et al. 2010), Amblyoponinae (Molet et al. 2007) and Ectatomminae (Lachaud et al. 1999b). On the other hand, small queens in E. tuberculatum colonies are inquiline parasites that have been described as a different species, Ectatomma parasiticum (Hora et al. 2005(Hora et al. , 2009Feitosa et al. 2008). Inquiline parasite queens lack worker progeny but coexist in the nest with the host queen(s), a type of permanent parasitism without slavery (Buschinger 2009). ...
... Clade 1 comprises three described species with a predominant distribution in South America: Ectatomma edentatum, E. muticum and Ectatomma suzanae. Clade 2 contains the wide-range E. tuberculatum, its recently described inquiline parasite, E. parasiticum (Feitosa et al. 2008), and Ectatomma vizottoi from Brazil. One of the remaining principal clades, clade 3, encompasses the other wide-range species E. ruidum, which is a sister to the Central American endemic Ectatomma gibbum. ...
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Os macrofungos têm sido pouco estudados nos diferentes biomas brasileiros e estima-se que há muitas espécies ainda a serem descritas. Este capítulo apresenta um levantamento incluindo a descrição de dezesseis espécies de Basidomycota (cogumelos lamelares e fungos clavarioides) coletados na Estação Ecológica do Rio Ronuro, Mato Grosso. Para cada espécie são apresentados dados taxonômicos e de distribuição geográfica no Brasil. Dez espécies são registradas pela primeira vez para o estado do Mato Grosso.
... Species of the genus Ectatomma are mainly predatory, hunting a variety of prey, but they also scavenge on dead arthropods or animals, and some species exploit sugary resources (fruit pulp, floral and extrafloral nectary, honeydew, or similar excretions) (Lachaud 2021). The genus, exclusively neotropical but widely distributed, contains few species: 14 extant and 1 fossil species currently recognized, and at least 4 cryptic species in the E. ruidum species complex, which, due to the presence of extensive mitochondrial heteroplasmy and the current impossibility of accurately distinguishing them on the basis of morphological characters, are still referred to as E. ruidum sp. 1, 2, 3, 4 (Kugler and Brown 1982;Feitosa et al. 2008;Nettel-Hernanz et al. 2015;Aguilar-Velasco et al. 2016;Meza-Lázaro et al. 2018Peña-Carrillo et al. 2021. Several of them, such as E. tuberculatum and E. ruidum sp. 1 and sp. 2, are considered dominant and important natural control agents in various agroecosystems (Perfecto 1990;Lachaud 1990Lachaud , 2021Majer et al. 1994;Ibarra-Núñez et al. 2001;Schatz and Lachaud 2008). ...
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Despite the extraordinary diversity of organisms associated with ants, few species or genera have been inventoried for the myrmecophilous communities they host. Here, we review the known information on Lepidoptera associated with the ant genus Ectatomma, based on: (a) extensive colony sampling and observations on six focal species (E. tuberculatum, E. brunneum, and four cryptic species of the E. ruidum species complex) over a period of 43 years in Mexico, French Guiana, and Colombia, (b) a follow-up of the available literature, and (c) an analysis of Ectatomma images from various photographic databases available online and direct observations in Brazil and Suriname. No lepidopteran species were found inside the nests, but a wide variety of facultative mutualisms were observed outside on plants bearing extrafloral nectaries and/or honeydew-producing hemipterans; however, around 15% involved a form of commensalism, with no direct physical butterfly–ant interaction. Various new associations, previously unnoticed, are reported, and we illustrate a new symbiotic association between Rekoa palegon and E. ruidum sp. 2 in Mexico. At least 29 lepidopteran species from 19 genera, belonging to four tribes in three subfamilies and three families, participate in 41 associations involving only 5 of the 18 known Ectatomma species, all 5 characterized by visiting liquid food sources on foliage. Specialized interactions with Ectatomma ants were only found in three Riodinidae species, while in Lycaenidae interactions were all facultative. A greater sampling effort is needed, including nocturnal sampling and studies on little-studied species of this genus, to obtain a comprehensive picture of the extent of Ectatomma–Lepidoptera interactions.
... Alternatively, these patterns have also been argued to be a product of historic sampling bias for ants in the northern hemisphere (Kutter, 1968;Wilson, 1971Wilson, , 1984. Fortunately, our knowledge about ant biodiversity in tropical regions has made significant progress in recent decades (see for example : Delabie et al., 2015;Fernández, 2003;Fernández et al., 2019;Fisher & Bolton, 2016;Shattuck, 2000), and simultaneously numerous ant social parasite species were discovered in tropical and subtropical regions (Bharti, Radchenko, et al., 2016;Blaimer, 2012a;De Souza et al., 2007;Dubovikoff & Longino, 2004;Feitosa et al., 2008;Fischer et al., 2020;Guerrero et al., 2010;Longino, 2003Longino, , 2006Longino, , 2007 Bacci, 2010;Rabeling et al., 2015Rabeling et al., , 2019Schultz et al., 1998). In fact, reviewing taxonomic studies including ant social parasites showed that the majority of newly discovered ant social parasites in the past two decades were described from the Neotropics (Appendix S1, Fig. S1.1). ...
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Aim One of the most consistent global biogeographic patterns is the latitudinal diversity gradient where species richness peaks within the equatorial tropics and decreases towards the poles. Here, we explore the global biogeography of socially parasitic ant species, which comprises the most diverse group of social parasites in the Hymenoptera. We test the biogeographic hypothesis that ant social parasites are distributed along an inverse latitudinal diversity gradient (iLDG) by peaking in diversity outside of the equatorial tropics, which would contrast with the biogeographic pattern observed in free‐living, non‐parasitic ant species. Location Global. Taxon Ants (Hymenoptera: Formicidae). Methods We assembled a comprehensive biogeographic dataset consisting of 6001 geographic distribution records for all 371 taxonomically described socially parasitic ant species. We used phylogenetic and taxonomic studies to estimate the number of independent evolutionary origins of ant social parasitism to directly compare species richness with the number of species representing independent evolutionary origins of social parasitism across a latitudinal gradient. In addition, we compared ant social parasite diversity across biogeographic regions using rarefaction to account for different sampling efforts. Finally, we tested for a correlation between latitude and the proportion of ant social parasite species within regional ant faunae. Results The geographic distribution records and the inferred 91 independent evolutionary origins of socially parasitic life histories in ants show that both species richness and the number of species representing independent evolutionary origins of social parasitism peak in the northern hemisphere outside of the equatorial tropics. Based on rarefaction curves, northern latitude regions harbour the most ant social parasite species, but the diversity of independent evolutionary origins is not significantly different between northern and southern hemispheres. The proportion of ant social parasite species within regional faunae is tightly correlated with latitude only in the northern hemisphere. Main conclusions The iLDG of ant social parasites contrasts with the biogeographic pattern observed in free‐living, non‐parasitic ant species and appears to be driven by large species radiations as well as by the presence of specialized life histories exclusive to the northern hemisphere.
... Small queen morphs, referred to as 'microgynes' , are in many species associated with alternative dispersal strategies and polygyny itself, and in some cases microgynes were shown to favor the production of sexual offspring over workers, providing the basis for reproductive cheating 30 . In some cases, microgynes are in fact considered intra-specific inquiline parasites 66,77 , whereas in other cases microgynous forms were raised to the species level as obligate inter-specific inquiline parasites 78,79 . Queen size polymorphism is heavily associated with the evolution of social parasitism based on two major lines of evidence. ...
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Social parasites exploit the brood care behavior of their hosts to raise their own offspring. Social parasites are common among eusocial Hymenoptera and exhibit a wide range of distinct life history traits in ants, bees, and wasps. In ants, obligate inquiline social parasites are workerless (or nearly-so) species that engage in lifelong interactions with their hosts, taking advantage of the existing host worker forces to reproduce and exploit host colonies’ resources. Inquiline social parasites are phylogenetically diverse with approximately 100 known species that evolved at least 40 times independently in ants. Importantly, ant inquilines tend to be closely related to their hosts, an observation referred to as ‘Emery’s Rule’. Polygyny, the presence of multiple egg-laying queens, was repeatedly suggested to be associated with the origin of inquiline social parasitism, either by providing the opportunity for reproductive cheating, thereby facilitating the origin of social parasite species, and/or by making polygynous species more vulnerable to social parasitism via the acceptance of additional egg-laying queens in their colonies. Although the association between host polygyny and the evolution of social parasitism has been repeatedly discussed in the literature, it has not been statistically tested in a phylogenetic framework across the ants. Here, we conduct a meta-analysis of ant social structure and social parasitism, testing for an association between polygyny and inquiline social parasitism with a phylogenetic correction for independent evolutionary events. We find an imperfect but significant over-representation of polygynous species among hosts of inquiline social parasites, suggesting that while polygyny is not required for the maintenance of inquiline social parasitism, it (or factors associated with it) may favor the origin of socially parasitic behavior. Our results are consistent with an intra-specific origin model for the evolution of inquiline social parasites by sympatric speciation but cannot exclude the alternative, inter-specific allopatric speciation model. The diversity of social parasite behaviors and host colony structures further supports the notion that inquiline social parasites evolved in parallel across unrelated ant genera in the formicoid clade via independent evolutionary pathways.
... Currently, the most comprehensive work including an identification key for the species in the genus is the revision by Kugler and Brown (1982). However, this work does not include the species Ectatomma parasiticum Feitosa and Fresneau, in Feitosa et al. (2008), E. suzannae Almeida (1986) andE. vizottoi Almeida (1987). ...
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Uncovering the evolutionary history of the subfamilies Ectatomminae and Heteroponerinae, or ectaheteromorphs, is key to understanding a major branch of the ant tree of life. Despite their diversity and ecological importance, phylogenetic relationships in the group have not been well explored. One particularly suitable tool for resolving phylogeny is the use of ultraconserved elements (UCEs), which have been shown to be ideal markers at a variety of evolutionary time scales. In the present study, we enriched and sequenced 2,127 UCEs from 135 specimens of ectaheteromorph ants and investigated phylogeny using a variety of model-based phylogenomic methods. Trees recovered from partitioned maximum-likelihood and species-tree analyses were well resolved and largely congruent. The results are consistent with an expanded concept of Ectatomminae that now includes the subfamily Heteroponerinae new synonym and its single tribe Heteroponerini new combination. Eleven monophyletic groups are recognized as genera: Acanthoponera, Alfariastatus revived, Boltonia Camacho and Feitosa new genus, Ectatomma, Gnamptogenys, Heteroponera, Holcoponerastatus revived, Poneracanthastatus revived, Rhytidoponera, Stictoponerastatus revived, and Typhlomyrmex. The new phylogenetic framework and classification proposed here will shed light on the study of Ectatomminae taxonomy and systematics, as well as on the morphological evolution of the groups that it comprises.
... In ants, social parasitism has been reported in six subfamilies: Dolichoderinae, Formicinae, Myrmeciinae, Pseudomyrmecinae, Myrmicinae (reviewed by Tinaut & Ruano, 1999) and Ectatomminae (Hora et al., 2005;Feitosa et al., 2008), among which, inquilinism is considered the most frequent type (Buschinger, 2009). In inquilinism sensu Wilson (1971), parasitic species spend their whole life cycle in the host's nest and frequently do not produce a worker caste, but instead invest resources and effort in producing only gynes and males. ...
Article
The ant Acromyrmex ameliae is a social parasite of two leaf-cutting ant subspecies: Acromyrmex subterraneus subterraneus and A. subterraneus brunneus. Cytogenetic data are available for 14 species of Acromyrmex and all of them possess 2n = 38 chromosomes. In this study, chromosome number, heterochromatin detection, and detection of AT and GC-rich blocks of colonies of A. ameliae and its hosts were carried out. Additionally, the detection of nucleolus organizer regions and 18S rDNA clusters in chromosomes of the parasite and physical mapping of telomeres were undertaken. The same chromosome number and morphology were detected for the hosts 2n = 38 (10m + 14sm + 12st + 2a), while the females and males of the social parasite A. ameliae presented 2n = 36 (10m + 16sm + 8st + 2a) and n = 18 (5m + 8sm + 4st + 1a). In both A. ameliae and its hosts, the terminal region on the short arm of the largest subtelocentric pair is heterochromatic GC-rich, and this region corresponded to the 18S rDNA clusters in the parasite. The short arms of several chromosomes were heterochromatin-rich. The telomeric probe hybridized telomeres on all chromosomes of the parasite and was not detected in intrachromosomal regions. Through a comparative cytogenetic analysis, we hypothesize that the karyotype of A. ameliae (2n = 36) originated from a chromosomal rearrangement that reduced the number of chromosomes from 38 to 36; as available data on the genus Acromyrmex show that all other species possess 38 chromosomes, representing 45% of the 33 valid species in this genus. The mechanism of the chromosome rearrangement is discussed. Thus, the chromosome number observed in A. ameliae is a derivation from the genus. Our data show variation in the chromosomal number in Acromyrmex and suggest that analyses of the karyotypes of parasite species can yield novel insights with regards to the evolution of this genus.
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Queen dimorphism is known in different ant subfamilíes and corresponds to the co-occurrence of severa! queen morphs within the same species. lt is frequently associated with different colony-founding and dispersion strategies. We studied the queen-size dimorphism in the Pon erina e ant, Ectatomma tuberculatum from field colonies collected at Apazapan (Veracruz, Mexico). Microgynes were present in half the colonies collected and their production among the female sexuals prevailed over that of macrogynes. Colonies possessed one or several gynes, up to 8 for macrogynes and 11 for microgynes. From the data basis on nest excavation, worker size population was correlated neither with the number of dealate gynes nor with the number of microgynes, and it seemed that microgynes were as productive as macrogynes. Both gynes were morphologically similar but their distribution was clearly bimodal without overlapping, the microgynes b eing not completely separated from the workers. Microgynes fell on a continuous regression line with the macrogynes and represented the isometric reduction of normal queens. The ovarían morphology differed among the three female groups: macrogynes had the most numerous ovarioles on average but produced as many oocytes as microgynes. Contrary to the workers, all gynes possessed a developed spermatheca and had the potential ability to lay fertilized eggs. Such queen dimorphism was not previously observed in this species and only found in the A paza pan si te. It may constitute an alternative reproductive strategy leading to a functional facultative polygyny. as already reported in E. ruidum (Schatz et al. 1996, Lachaud et al. l 999a, b). or a case of a social intraspecific parasitism. Further investigations are being conducted to better understand the reproductive biology of this species.
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Ectatomma tuberculatum (Olivier) preys primarily on small invertebrates, including other social insects. In Panama, workers forage primarily at night and nocturnal activity begins with a mass exit at dusk. Several times a day, particularly at dawn and dusk, hundreds of Crematogaster limata Fr. Smith file into nest entrances of E. tuberculatum. These cleptobiotic encounters may represent a primitive form of social parasitism.
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This study provides a detailed account of the foraging behaviour of the ponerine ant Ectatomma opaciventre in a ‘cerrado’ savannah in south-east Brazil. Our observations suggest that this species has an exclusively diurnal foraging pattern. Feeding habits included both predation and scavenging, with termite workers and leaf-cutting ants as the most important food items. Contrary to all other Ectatomma species studied to date, no liquid food such as hemipteran honeydew or plant nectar was collected. Foragers showed clear individual foraging area fidelity.Workers of E. opaciventre employed a typical individual foraging strategy, i.e. there was no co-operation between foragers in the search for or retrieval of food, neither by tandem running nor by trail laying. Nest density was considerably lower than in other Ectatomma (0.015 nests per m2). The observed mean distance to the nearest neighbouring nest was 5.85 m, with a significant tendency toward over-dispersion. Nests were more frequently found in specific microhabitats, which may suggest active choice of nesting site by founding queens.
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In the ponerine ant Ectatomma tuberculatum, workers display an ovariole cycle composed of nine distinct stages. Ovarian maturation depends partly on age. During the fist 4 weeks of imaginal life, the ovarioles develop a succession of polytrophic follicles. They reach their maximum growth at approximately 45 days old; at about 3 months, they begin a period of regression until they are completely inactivated at the age of approximately 1 year. However, workers of exactly the same age may display up to five different stages. Such variability is not related to differences in worker size. Body size seems to affect only the number of ovarioles (and perhaps the ability to produce alimentary eggs).
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A taxonomic review is made of the 15 described species of socially parasitic Myrmica, found in the Palaearctic, and 3 apparently free-living Myrmica species that have characteristics of the "parasitic syndrome". Notes on the current taxonomic status and biological knowledge of each species are given. Earlier synonymies are discussed and one new synonymy is made: M. samnitica Mei = M. laurae Emery. Also, the synonymy of M. myrmecophila Wasmann with M. sulcinodis Nylander is confirmed and it is suggested that the type specimen is neither an ergatoid queen nor a social parasite, but a worker parasitized by Mermis. The status of M. symbiotica Menozzi remains unclear: it is not an ergatoid queen but could be a pseudogyne worker of a parasitic species with as yet undescribed queens. Keys are given for the identification of all castes of the 10 recognised species of social parasite (including M. symbiotica) and the 3 associated free-living species.