# Diapriinae Wasps (Hymenoptera: Diaprioidea: Diapriidae) Associated with Ants (Hymenoptera: Formicidae) in Argentina

Authors: Marta S. Loiácono; Cecilia B. Margaría; Daniel A. Aquino
Journal: Psyche
 (2013)

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## Abstract

We provide an overview of diapriid wasps associated with ants in Argentina and the diversity of interactions they have developed with their hosts. As a result, we report 16 species of nine genera of Diapriinae, two new geographic distributions, three new association records, illustrations, and photographs. We highlight myrmecophile symphylic species, with a high degree of integration with the host ants, adaptation being morphological and behavioral. A table with diapriid species and ant hosts is given.


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## Body

## 1. Introduction

Diapriids are primary endoparasitoids of larvae-pupae or pupae, principally of dipterans, but a number of species are closely associated with ant nests. However, there are few behavioral data on host-diapriid myrmecophile interactions. Huggert and Masner [1] hypothesized that the ancestors of diapriines guests changed from Diptera to Formicidae. The intermediates in the presumed sequence of hosts seem to be the numerous synoeketic Diptera living in the refuse depot  and bivouacs of various army ants of the subfamily Ecitoninae. Diapriines females, in the search for potential hosts, would have progressively integrated with formicids. According to Masner (personal communication) this change would have occurred more frequently in the Neotropical region where these ants have high distribution. The guests switch mechanism has determined morphological and behavioral specialization, manifested by the degree of integration of diapriines to ant colonies. These symphyles are often highly adapted to their hosts, exhibiting morphological and behavioral adaptations to living with ants (extensive morphological mimicry of the host ants coloration, ocellus regression, similar sculpture, presence of appeasement  substances in specialized structures and trichomes, trophallaxis, etc.), which aid them in avoiding detection and/or aggression by host ants. Ants seem to have preference to lick certain parts of diapriid body to get exudates [2]. The adaptations include secondary apterism in which the wings of wasps are bitten off by either the parasite itself or its host. During the alate phase, the adults probably disperse, as the alate individuals, caught by sweeping, in Malaise traps and significantly by light traps indicating also the nocturnal activity in this phase of life [2]. The secondary apterism occurs in several species of diapriines, for example, Asolenopsia rufaKieffer, Bruchopria pentatomaKieffer, Bruchopria hexatoma Kieffer, Notoxoides pronotalis (Borgmeier), herein studied.The current knowledge indicates that only a few diapriids are parasitoids of ant brood, attacking as solitary or gregarious koinobiont endoparasitoids of the host larvae, and worker and/or reproductive immature stages can be parasitized. From 121 diapriine species in 34 genera that had been collected in association with ants, development of immature stages as parasitoids of ant larvae has been demonstrated for only 26 species in seven genera, most of which are only known at the level of morphospecies [3]. There are only two species and one morphospecies recorded in Argentina as ant parasitoids [4].A large number of diapriine wasps became associated with various groups of ants in Central and South America. The associations are especially well developed with army ants (Ecitonini) and leaf cutting ants (Attini) with some 20 genera of Diapriinae already involved [5]. The vast majority of these species belong to Diapriini, although there are some exceptions like Bruchopriaspecies that belong to the tribe Spilomicrini [6].The New World fungus-growing ants (Hymenoptera: Formicidae: Attini) are especially diverse in the tropics. As true for the most social insects, they accumulate significant stores of resources within their nests, attracting a diverse array of predators, microbial pathogens, and parasites [7]. We studied aspects of the intensity and prevalence of these little-known diapriine wasps that attack the larvae of the fungus-growing ant, Acromyrmex lobicornisEmery, and noted a remarkably diverse community of parasitoids within host population from four localities of La Pampa, Argentina [4, 8]. In some cases, the rates of parasitoidism can reach high levels. Loiácono et al. [4] collected 1560 wasps (adults and immatures) from 430 parasitized larvae from three partial colonies of Acromyrmex, which shows how prevalent these wasps can be in attacking the ants. Fernández-Marin et al. [9] found that between 27% and 70% of the colonies of two species of Cyphomyrmex Mayr  were parasitized by one species in Puerto Rico and by up to four concurrent morphospecies of diapriids in Panama. Similarly, Pérez-Ortega et al. [7] reported that another fungus-growing ant, Trachymyrmex cf.zeteki, was attacked by a diverse community of diapriids in Panama, with a mean intensity of larval parasitism per ant colony of 33.9%, and prevalence across all ant populations of 27.2%. Lachaud and Pérez Lachaud [3], based on the abundance and success in attacking ants, considered that diapriids and another group of microhymenopterans, the eucharitids, seem excellent potential models to explore how parasitoids impact ant colony demography, population biology, and ant community structure [3].In Argentina, the study of myrmecophiles has attracted the attention of several scientists in the last two centuries. Carlos Bruch (1869–1943), a German naturalist selected by F. Moreno—first Director of Museo de La Plata—to organize its collections, was a pioneer of the entomological studies; it is important to remark his ability as a photographer and scientific illustrator, and his observations regarding special associations and behaviors of ants and beetles: termitophily and myrmecophily [10, 11]. Jean-Jacques Kieffer (1857–1925), a French entomologist who specialized in the study of parasitoids of insects, based his studies on Bruch’s material and published articles about diapriines associated with ants [12, 13]. Alejandro Ogloblin (1891–1967), a Russian entomologist researcher at “Estación Experimental de Loreto” (Misiones, Argentina), collected there numerous diapriid wasps associated specially with myrmicine ants [14, 15]. Luis De Santis (1914–2000) catalogued associations between diapriids and ants [16, 17] and reported new geographic distributions [18]. Marta Loiácono and colleagues studied Neotropical myrmecophiles diapriids and their interactions with ants [4, 7, 8, 15, 19–29].In this paper, we provide an overview of the diversity of diapriid wasps associated with ants in Argentina and the diversity of interactions they have developed with their hosts.


## 2. Material and Methods

Specimens for this study were reared in laboratory [4] or collected from ant nests, killed in alcohol, and mounted on cards or microscopic slides for further studies. Observations of the specimens were made through a stereomicroscope Leica S8APO. The photographs were taken by Daniel A. Aquino with a Leica DFC295 camera attached to the stereomicroscope. Digital images were mounted using open software CombineZM [30] and enhanced using Photoshop. Scanning micrographs were taken with a JEOL JSMT100 at Museo de La Plata operating at 15 KV.Sharkey [31] was followed for the higher-level phylogeny of the Hymenoptera order, Bolton for ant valid names [32], Masner and García [5] for diapriid systematics, and Yoder et al. web site [33] for interactive keys and links.Diapriid and ant specimens examined in this study are deposited at Museo de La Plata (Buenos Aires, Argentina). Most of them were collected and determined by Bruch and Ogloblin in Argentina. Type material ofSzelenyiopria reinchenspergeri (Ferrière) was loan by Hungarian Natural History Museum.Biology Section includes “hosts” wasps emerged from ant larvae or “associated” wasps found in or near nests or emigration columns of army ants.


## 3. Results

### 3.1. Tribe Diapriini Ashmead, 1893 [34]

#### 3.1.1.Asolenopsia Kieffer, 1921 [12]

Asolenopsia Kieffer, 1921: 36 [12].Euplacopria Ferrière, 1929: 157 [35].Distribution
Tropical lowlands of Central and South America [5].Biology
Associated with ecitonini ants of genusEciton Latreille, Labidus Jurine  and Neivamyrmex Borgmeier [5].Remarks
Members ofAsolenopsia are moderately to highly specialized associates to ecitonine ants [20]. Their wings are primarily developed but subsequently bitten off by ants or cast off spontaneously (alectomy). Winged adults are also collected in light traps [5].


#### 3.1.2.Asolenopsia rufa Kieffer, 1921 [12] (Figure 1(a))

(a)Asolenopsia rufa female dealate in dorsal view. (b) Neivamyrmex carettei.Scale: 1 mm.
(a)
(b)Asolenopsia rufa Kieffer, 1921: 37 [12].Distribution
Argentina (Córdoba, Entre Ríos, and Santa Fe) [12, 17].Biology
Associated withNeivamyrmex carettei (Forel) [12] (Figure 1(b)).Material Studied
Syntype, female, dealated, withNeivamyrmex carettei worker, Argentina, Córdoba, Alta Gracia, La Granja, 1-8-IV-1920, Bruch coll.; one female, without date, Santa Fe, Vera y Pintado (Fives Lille), Weiser coll.; female alated, Argentina, Misiones, Loreto, without date, Ogloblin coll.


#### 3.1.3.Basalys Westwood, 1832 [36]

BasalysWestwood, 1832: 342–344 [36].CeratopriaAshmead, 1893: 407, 42 [34].Acidopria Kieffer, 1913: 442 [37].Loxotropa auct. nec Foerster, synonymized by Masner, 1964 [38].Nesopria Muesebeck and Walkley, 1956: 319–419 [39].Distribution
The genus is well represented in North and South America, rarely in Chile [5].Biology
Several species were reared from various dipterous hosts, and some were collected in ant nests [5].


#### 3.1.4.Basalys sp.

Material Studied
One female and 1 male (microscopic slide) collected with the “Argentine ant,”Linepithema humile (Mayr), Argentina, Buenos Aires, J. C. Paz, 11-X-1934, Ogloblin coll.; 1 female (microscopic slide) collected with the Argentine ant Linepithema humile, Argentina, Buenos Aires, J. C. Paz, 8-IX-1945, Bezzi leg.Biology
Associated withLinepithema humile (new record).Remarks
Female and male studied were determined by Masner, who wrote a label: “Basalys sp. ♀♂(=Loxotropa auct.) aberrant sp. with !11-segm. ant. ♀, Det. L. Masner, ‘89”; and female specimen: “Basalys sp. ♀(=Loxotropa auct.) !11-segmented antenna, Det. L. Masner, ’89.” Specimens studied were determined by Ogloblin as a new species of Doliopria, but he did not describe it. We also considered that material studied belong to genus Basalys, as it was established by Masner.


#### 3.1.5.Doliopria Kieffer, 1910 [40]

Doliopria Kieffer, 1910: 48 [40].Martinica Risbec, 1950: 533 [41].Distribution
Doliopria is restricted to the New World, with only a few species in the Nearctic region and with a high number of undescribed species in tropical America [5].Biology
Associated with ecitonini and attini ants [5].Remarks
Three Neotropical species were described associated with ants [12, 35, 40]; hypothetically they parasitized synoeketic Diptera because they show no specialized morphology [5].


#### 3.1.6.Doliopria collegii Ferrière, 1929 [35] (Figure 2)

Figure 2
Doliopria collegii female in lateral view. Scale: 1 mm.Doliopria collegii Ferrière, 1929 : 164 [35].Distribution
Argentina (Buenos Aires and Misiones) [18, 35].Biology
Associated with ecitonini ants,Eciton burchellii (Westwood) and Eciton quadriglume (Haliday) [35].Material Studied
Two females alated, Argentina, Misiones, Loreto, 20-X-1919 and 18-IX-1923, Ogloblin coll. and det.


#### 3.1.7.Doliopria myrmecobia Kieffer, 1921 [12] (Figure 3(a))

(a)Doliopria myrmecobia female in lateral view. (b) Acromyrmex lundii.Scale: 1 mm.
(a)
(b)Doliopria myrmecobia Kieffer, 1921: 39 [12].Distribution
Argentina (Buenos Aires; Misiones, new record) [12].Biology
Associated with attini antsAcromyrmex lundii (Guérin-Méneville) [12] (Figure 3(b)).Material Studied
One female, Argentina, Buenos Aires, La Plata, VIII, inside a nest ofAcromyrmex lundii, Bruch coll.; 1 female, alated collected with Acromyrmex sp., Argentina, Misiones, Loreto, 3-XI-1928, Ogloblin coll. and det.


#### 3.1.8.Notoxoides Ashmead, 1903 [42]

Notoxoides Ashmead, 1903: 30 [42].NotoxopriaKieffer, 1910: 39 [40].Philolestes Kieffer, 1922: 205 [13].Psilogasteroides Brèthes, 1911: 209–210 [43].Distribution
Restricted to lowland rainforests of continental South America [5].Biology
Members ofNotoxoides display some of the most advanced associations with ants. So far, ants of genera Neivamyrmex and Eciton (Ecitonini) were recorded as hosts [19]. Adult wasps are frequently collected in light traps. Wings may be lost to typical alectomy as indicated by shriveled wing rudiments in some specimens [5].


#### 3.1.9.Notoxoides pedissequus (Borgmeier, 1939) [44]

Notoxopria pedissequaBorgmeier, 1939: 538 [44].Distribution
Argentina (Córdoba) [19].Biology
Associated withNeivamyrmex pseudops (Forel) [44].Remarks
Loiácono [20] studied a female alate collected by Bruch in Córdoba province.


#### 3.1.10.Notoxoides pronotalis(Borgmeier, 1939) [44] (Figures 4(a), 4(b), and 5)

Notoxoides pronotalis female in dorsal view: (a) alate and (b) dealate specimens. Scale: 1 mm. (c) Eciton dulciumcollected with Notoxoides pronotalis, in lateral view.Scale: 1 mm.
(a)
(b)
(c)Figure 5
Original illustration ofNotoxoides pronotalisfemale in lateral view, by Bruch.Philolestes rufus Kieffer, 1922: 205 [13].Philolestes pronotalis Borgmeier, 1939: 536 [44].Notoxoides pronotalis: Masner, 1977: 34 [45].Notoxoides kiefferi Loiácono, 1981: 305, 306 [19].Distribution
Argentina (Córdoba, Salta, San Luis, and Santiago del Estero) [19, 44].Biology
Associated with Eciton dulcium Forel andNeivamyrmex sulcatus (Mayr) [44].Material Studied
Syntype, female dealated, collected withEciton dulcium, Argentina, Córdoba, Alta Gracia, 4-XII-I921, Bruch coll.; 2 syntype females alated, same data as syntype except II-1922, collected with Neivamyrmex sulcatus, Bruch coll. and det.; 21 females dealated, Argentina, Salta, Tartagal, I-1960, Martínez coll., with a Eciton dulcium, and 5 females alated, Argentina, Salta, Pocitos, III-1959, Martínez coll.; 3 females dealated and 1 alated, Córdoba, San Javier, La Paz, 15-31-XII-1928, Bruch coll., with Eciton dulcium; Córdoba, Alta Gracia: 1 female dealated, collected with Eciton dulcium (Figure 4(c)), 4-XII-1922, Bruch coll.; 1 female dealated,  without date and collector; 1 female alated, La Granja, 21-VIII-1924, Bruch coll.; 2 females dealated, La Granja, 25-I-1925, Bruch coll.; 3 females alated, La Granja, 4-XI-1925, Bruch coll.; 2 females dealated, with Eciton dulcium, La Granja, 4-XI-1925, Bruch coll.; 2 females alated with Eciton dulcium, 13-III-1934, Bruch coll.;  3 females alated, without date, Bruch coll.; 1 female dealated; Córdoba, Unquillo, without date and collector; 9 females alated, Córdoba, Unquillo, without date and collector; 1 female with fore wings, Córdoba, Unquillo, without date and collector; 2 females alated, Santiago del Estero, Cerrillos, 2-V-1955, without collector, and 2 females alated, without date, Bruch coll.; 5 females alated, without locality, 21-II-1925, light collected, without collector; 2 females dealated and 3 alated, without locality, 22-II-1925, light collected, without collector; 5 females alated and 1 dealated, without locality, 23-II-1925, light collected, without collector; 1 female alated, without locality, 24-II-1925, light collected, without collector.Remarks
Bruch always sent to Kieffer diapriid samples to be studied. As we mentioned, he was an excellent scientific illustrator (Figure5) [46] and an important photographer as is shown in (Figure 6) Neivamyrmex pseudops, ant host of Notoxoides pedisequus [47].Figure 6
Neivamyrmex pseudops,gravid queen in dorsal view, photographed by Bruch.We observed numerous both alate and dealate individuals found dependent on the phase of life. As is mentioned [2], during the alate phase, numerous adults were caught by light traps as we observed in the female material light collected by Bruch.Lachaud [48] mentioned that ants search actively for some chemical substances produced by glands at the basis of the setae present on the diapriid cuticle; similarly we observed the presence of peculiar neck hairs in N. pronotalis [20].


#### 3.1.11.Szelenyiopria Fabritius, 1974 [49]

SzelenyiopriaFabritius, 1974: 54 [49].Gymnopria Loiácono, 1987: 130 [21].Distribution
Wide distribution from Argentina to Guatemala [21, 49].Biology
Szelenyiopria lucens (Loiácono) from Uruguay is the first member of the tribe Diapriini in the New World positively reared from ants. Loiácono [21] reports up to three wasps per mature larva of Acromyrmex ambiguus (Emery) (Attini). Members of Szelenyiopria show no specialized structures known among other myrmecophilic Diapriini; Masner and García [5] assumed that the specialized setae with truncate apices are outlet of chemical substances.


#### 3.1.12.Szelenyiopria pampeana (Loiácono, 2000) [4] (Figure 7(a))

(a)Szelenyiopria pampeanafemale in lateral view. (b) Acromyrmex lobicornis larva showing immature instars of diapriines.Scale: 1 mm.
(a)
(b)Gymnopria pampeana Loiácono, 2000: 10 in Loiácono et al., 2000 [4].Szelenyiopria pampeana: Loiácono and Margaría, 2009: 63 [8].Distribution
Argentina (La Pampa) [4, 8].Biology
Koinobiont and gregarious endoparasitoids of late instar larvae ofAcromyrmex lobicornis (Emery), it was also established simultaneous parasitoidism with Trichopria sp. [4] (Figure 7(b)).Material Studied
Holotype female, Argentina, Santa Rosa, 8-XI-1995, Quirán and Corró Molas colls.; 25 paratypes females and 3 males, Lihuel Calel, 4-XII-1997,  Quirán and Corró Molas colls.


#### 3.1.13.Szelenyiopria reichenspergeri (Ferrière, 1929) [35]

Doliopria reinchespergeri Ferrière, 1929: 165 [35].Szelenyiopria reinchespergeri: Fabritius, 1974, 54 [49].Distribution
Argentina (Salta and Tucumán) [35, 49].Biology
Associated withEciton quadriglume and Neivamyrmex legionis (Smith) [35, 49].Material Studied
One female, Argentina, Salta, 2-6-II-1950, Golbach coll.


#### 3.1.14.Szelenyiopria sp.

Distribution
Argentina (Córdoba) (new record).Material Studied
Female and 3 males with an ecitonine ant, Argentina, Córdoba, San Javier, La Paz, 1-20-I-1929, Bruch coll.Remarks
Most females of this genus have 11-segmented antennae, but material studied here presents antenna 12-segmented as mentioned by Masner and García [5] for undescribed species. We considered that these specimens belong to Szelenyiopria genus by the most important feature, the presence on entire body of specialized straight setae, truncate apically.


#### 3.1.15.Trichopria Ashmead, 1893 [34]

Trichopria Ashmead, 1893: 407, 431 [34].Ashmeadopria Kieffer, 1912: 8, 10, 59 [50].Phaenopria Ashmead, 1893: 40, 436 [34].Planopria Kieffer, 1906: 19 [51].Orthopria Kieffer, 1911: 983, 984 [52].Distribution
Worldwide [5].Biology
Associated with the “fire ant,”Solenopsis richteri Forel (Kieffer, 1921) and endoparasitoid of Acromyrmex lobicornis [4].


#### 3.1.16.Trichopria formicans Loiácono, 2000 [4] (Figures 8(a) and 8(b))

Trichopria formicans female (a) in dorsal view and (b) lateral view. Scale: 1 mm.
(a)
(b)Trichopria formicans  Loiácono 2000 in Loiácono et al., 2000: 12 [4].Distribution
Argentina (La Pampa) [4].Biology
Reared from larvae ofAcromyrmex lobicornis [4].Material Studied
Holotype female, Argentina, La Pampa, Utracán, 22-XII-1997, Caramuti y Rodriguez colls.; paratypes 68 females and 43 males (MLP), same data as holotype.


#### 3.1.17.Trichopria myrmecophila (Kieffer, 1921) [12]

Phaenopria myrmecophila Kieffer, 1921: 4 [12].Trichopria myrmecophila: De Santis in De Santis and Esquivel, 1966: 50 [16].Distribution
Argentina (Buenos Aires) [12].Biology
Associated withSolenopsis richteri [12].


#### 3.1.18.Trichopria sp.

Distribution
Argentina Buenos Aires.Biology
Collected with the “argentine ant,”Linepithema humile (new record).Material Studied
Female collected withLinepithema humile, Argentina, Buenos Aires, J. C. Paz, 8-II-1940, Ogloblin coll.Remarks
Masner studied this material and determined specimens asTrichopria s. str. sp.


### 3.2. Tribe Spilomicrini Ashmead, 1893 [34]

#### 3.2.1.BruchopriaKieffer, 1921 [12]

Bruchopria Kieffer, 1921: 38 [12].Aulatopria Brèthes, 1927: 164 [53].Distribution
Argentina (Buenos Aires, Córdoba, and Misiones) [12, 53].Biology
Associated with ants of the generaSolenopsis Westwood (Solenopsidini) and Acromyrmex  Mayr (Attini) [12].Remarks
Hölldobler and Wilson [54] mentioned specimens of genus Bruchopria, as Solenopsis guest. Masner and García [5] mentioned “wings often bitten off by ants.” Loiácono et al. [26] studied alated and dealated individuals of Bruchopria species. The action of dealation has not been observed. The presence of tegulae with normal development and wing stumps demonstrates that the apterism has a secondary origin, caused by the autotomy or by bites of the host ants. The apices of the wing stumps of all individuals examined were regular suggesting that the wings are bitten or torn off close to the tegulae. The fact that specimens are dealated allows them to move into the mound galleries and chambers.


#### 3.2.2.Bruchopria hexatoma Kieffer, 1921 [12] (Figures 9(a), 10(a), and 10(b))

(a)Bruchopria hexatoma female dealate in dorsal view. (b) Solenopsis richteri.Scale: 1 mm.
(a)
(b)Bruchopria hexatoma female. (a) mesosoma and petiole in dorsal view, scale: 0.5 mm; (b) wing stump, scale: 0.1 mm [26].
(a)
(b)Bruchopria hexatoma
Kieffer, 1921: 39 [12].Bruchopria hexatoma: Borgmeier, 1939: 543 [44].Distribution
Argentina (Misiones, Córdoba and Buenos Aires) [12, 44].Biology
Associated withSolenopsis richteri (Figure 9(b)) and Acromyrmex lundii [12, 44].Material Studied
One female dealated, Argentina, Misiones, Pastoreo Grande, 9-VII-1932, Ogloblin coll.; 1 female dealated, Argentina, Córdoba, XII-1920, Bruch coll., 1 female dealated, Córdoba, Sierras de Córdoba, La Granja, Bruch coll., without date; 1 male dealated, Argentina, Buenos Aires, without locality, 9-VII-1923, Bruch coll., with the ant; 4 females dealated, Argentina, Buenos Aires, Olivos, without date, Bruch coll., with the ant; 1 female dealated, Argentina, Buenos Aires, 10-IX-1925, Bruch coll.; 1 female dealated withAcromyrmex lundii, Argentina, Buenos Aires, without date, Bruch coll.Remarks
Bruchopria hexatoma has been reported by Kieffer [12] in association with Solenopsis richteri and Acromyrmex lundii in Argentina; Borgmeier [44] also mentioned this species as a guest of S. saevissima (Smith), in Brazil.The specimens from the provinces of Córdoba and Buenos Aires are dealated, with remains of wings (Figures10(a) and 10(b)), and most of them are accompanied by the host ants (Figure 9(b)). Unfortunately, the types of the species described by Kieffer have become widely scattered or lost [55]. Bruch sent to Kieffer part of the same series of material to identify (De Santis, pers. comm.).


#### 3.2.3.Bruchopria pentatoma Kieffer, 1921 [12]

Bruchopria pentatoma Kieffer, 1921: 38 [12].Distribution
Argentina (Córdoba) [12].Biology
Associated withSolenopsis richteri [12].Material Studied
Syntype male dealated, Argentina, Córdoba, Alta Gracia; 1-8-IV-1920, Bruch coll.Remarks
According to Kieffer's description, females of both species,B. pentatoma and B. hexatoma, are distinguished by the number of club antennomeres, five and six, respectively. Unfortunately, the unique female type is not available. Bruchopria pentatoma has also been reported by Kieffer [12] in association with S. richteri and Acromyrmex lundii (Guérin) in Argentina.


#### 3.2.4.PentapriaKieffer, 1905 [56]

Pentapria Kieffer, 1905: 34 [56].Antipapria Fabritius, 1968: 844 [57].Bakeria Kieffer, 1905: 34 [56].Plutopria Kieffer, 1910: 48 [40].Spilomicrinus Ogloblin, 1957: 425 [58].Xenopria Fouts, 1939: 260 [59].Distribution
The genus is distributed in the New World [5].Biology
The principal host plausible to assume is Stratiomyidae (Diptera) [5]. Herein, we studied a female collected with Solenopsis saevissima (Hymenoptera: Formicidae).


#### 3.2.5.Pentapria cf. nodicornis

Distribution
Argentina (Córdoba).Biology
Associated withSolenopsis saevissima (new record).Material Studied
Female collected withSolenopsis saevissima, Argentina, Córdoba, Alta Gracia, La Granja, II-1927, Bruch. coll., with no more data.


#### 3.2.6.SpilomicrusWestwood, 1832 [36]

SpilomicrusWestwood, 1832: 129 [36].LoxotropaFoerster, 1856: 122, 123, 126 [60].HoplopriaAshmead, 1893: 385, 386, 388 [34].LinkiolaKieffer, 1910: 39 [40].EriopriaKieffer, 1910: 693, 744 [40].TritopriaKieffer, 1910: 717, 748 [40].CologlyptusCrawford, 1910: 123 [61].ScutellipriaSzabó, 1961: 53–493 [62].Distribution
America [5].Biology
Primary parasitoidism solitary and gregarious of various Diptera; few species were reared from Coleoptera [5]. Herein, we studied samples associated with a Solenopsidini ant.


#### 3.2.7.Spilomicrus sp.

Distribution
Argentina (Buenos Aires).Biology
Associated with Solenopsidini ant.Material Studied
Two females with a Solenopsidini ant, Argentina, Buenos Aires, 9-VIII-1923, Bruch coll.Table1 summarizes information about diapriids and their associates.Table 1
Diapriid tribe
Diapriids species
Argentine provinces
Ant subfamily
Ant tribe
Ant species
Diapriini
Asolenopsia rufa
Córdoba, Entre Ríos, Santa Fe
Ecitoninae
Ecitonini
Neivamyrmex carettei
Basalyssp.
Buenos Aires
Dolichoderinae
Dolichoderini
Linepithema humile
Doliopria collegii
Buenos Aires, Misiones
Ecitoninae
Ecitonini
Eciton burchellii, Eciton quadriglume
Doliopria myrmecobia
Buenos Aires, Misiones
Myrmicinae
Attini
Acromyrmex lundii
Notoxoides pedissequus
Córdoba
Ecitoninae
Ecitonini
Neivamyrmex pseudops
Notoxoides pronotalis
Córdoba, Salta, San Luis,  Santiago del Estero
Ecitoninae
Ecitonini
Eciton dulcium, Neivamyrmex sulcatus
Szelenyiopria pampeana
La Pampa
Myrmicinae
Attini
Acromyrmex lobicornis
Szelenyiopria reichenspergeri
Salta, Tucumán
Ecitoninae
Ecitonini
Eciton quadriglume, Neivamyrmex legionis
Szelenyiopriasp.
Córdoba
Ecitoninae
Ecitonini
Ecitonini sp.
Trichopria formicans
La Pampa
Myrmicinae
Attini
Acromyrmex lobicornis
Trichopria myrmecophila
Buenos Aires
Myrmicinae
Solenopsidini
Solenopsis richteri
Trichopria sp.
Buenos Aires
Dolichoderinae
Dolichoderini
Linepithema humile
Spilomicrini
Bruchopria hexatoma
Buenos Aires, Córdoba, Misiones
MyrmicinaeMyrmicinae
SolenopsidiniAttini
Solenopsis richteri
Acromyrmex lundii
Bruchopria pentatoma
Córdoba
Myrmicinae
Solenopsidini
Solenopsis richteri
Pentapria cf. nodicornis
Córdoba
Myrmicinae
Solenopsidini
Solenopsis saevissima
Spilomicrussp.
Buenos Aires
Myrmicinae
Solenopsidini
Solenopsidini sp.


## 3.1. Tribe Diapriini Ashmead, 1893 [34]

### 3.1.1.Asolenopsia Kieffer, 1921 [12]

Asolenopsia Kieffer, 1921: 36 [12].Euplacopria Ferrière, 1929: 157 [35].Distribution
Tropical lowlands of Central and South America [5].Biology
Associated with ecitonini ants of genusEciton Latreille, Labidus Jurine  and Neivamyrmex Borgmeier [5].Remarks
Members ofAsolenopsia are moderately to highly specialized associates to ecitonine ants [20]. Their wings are primarily developed but subsequently bitten off by ants or cast off spontaneously (alectomy). Winged adults are also collected in light traps [5].


### 3.1.2.Asolenopsia rufa Kieffer, 1921 [12] (Figure 1(a))

(a)Asolenopsia rufa female dealate in dorsal view. (b) Neivamyrmex carettei.Scale: 1 mm.
(a)
(b)Asolenopsia rufa Kieffer, 1921: 37 [12].Distribution
Argentina (Córdoba, Entre Ríos, and Santa Fe) [12, 17].Biology
Associated withNeivamyrmex carettei (Forel) [12] (Figure 1(b)).Material Studied
Syntype, female, dealated, withNeivamyrmex carettei worker, Argentina, Córdoba, Alta Gracia, La Granja, 1-8-IV-1920, Bruch coll.; one female, without date, Santa Fe, Vera y Pintado (Fives Lille), Weiser coll.; female alated, Argentina, Misiones, Loreto, without date, Ogloblin coll.


### 3.1.3.Basalys Westwood, 1832 [36]

BasalysWestwood, 1832: 342–344 [36].CeratopriaAshmead, 1893: 407, 42 [34].Acidopria Kieffer, 1913: 442 [37].Loxotropa auct. nec Foerster, synonymized by Masner, 1964 [38].Nesopria Muesebeck and Walkley, 1956: 319–419 [39].Distribution
The genus is well represented in North and South America, rarely in Chile [5].Biology
Several species were reared from various dipterous hosts, and some were collected in ant nests [5].


### 3.1.4.Basalys sp.

Material Studied
One female and 1 male (microscopic slide) collected with the “Argentine ant,”Linepithema humile (Mayr), Argentina, Buenos Aires, J. C. Paz, 11-X-1934, Ogloblin coll.; 1 female (microscopic slide) collected with the Argentine ant Linepithema humile, Argentina, Buenos Aires, J. C. Paz, 8-IX-1945, Bezzi leg.Biology
Associated withLinepithema humile (new record).Remarks
Female and male studied were determined by Masner, who wrote a label: “Basalys sp. ♀♂(=Loxotropa auct.) aberrant sp. with !11-segm. ant. ♀, Det. L. Masner, ‘89”; and female specimen: “Basalys sp. ♀(=Loxotropa auct.) !11-segmented antenna, Det. L. Masner, ’89.” Specimens studied were determined by Ogloblin as a new species of Doliopria, but he did not describe it. We also considered that material studied belong to genus Basalys, as it was established by Masner.


### 3.1.5.Doliopria Kieffer, 1910 [40]

Doliopria Kieffer, 1910: 48 [40].Martinica Risbec, 1950: 533 [41].Distribution
Doliopria is restricted to the New World, with only a few species in the Nearctic region and with a high number of undescribed species in tropical America [5].Biology
Associated with ecitonini and attini ants [5].Remarks
Three Neotropical species were described associated with ants [12, 35, 40]; hypothetically they parasitized synoeketic Diptera because they show no specialized morphology [5].


### 3.1.6.Doliopria collegii Ferrière, 1929 [35] (Figure 2)

Figure 2
Doliopria collegii female in lateral view. Scale: 1 mm.Doliopria collegii Ferrière, 1929 : 164 [35].Distribution
Argentina (Buenos Aires and Misiones) [18, 35].Biology
Associated with ecitonini ants,Eciton burchellii (Westwood) and Eciton quadriglume (Haliday) [35].Material Studied
Two females alated, Argentina, Misiones, Loreto, 20-X-1919 and 18-IX-1923, Ogloblin coll. and det.


### 3.1.7.Doliopria myrmecobia Kieffer, 1921 [12] (Figure 3(a))

(a)Doliopria myrmecobia female in lateral view. (b) Acromyrmex lundii.Scale: 1 mm.
(a)
(b)Doliopria myrmecobia Kieffer, 1921: 39 [12].Distribution
Argentina (Buenos Aires; Misiones, new record) [12].Biology
Associated with attini antsAcromyrmex lundii (Guérin-Méneville) [12] (Figure 3(b)).Material Studied
One female, Argentina, Buenos Aires, La Plata, VIII, inside a nest ofAcromyrmex lundii, Bruch coll.; 1 female, alated collected with Acromyrmex sp., Argentina, Misiones, Loreto, 3-XI-1928, Ogloblin coll. and det.


### 3.1.8.Notoxoides Ashmead, 1903 [42]

Notoxoides Ashmead, 1903: 30 [42].NotoxopriaKieffer, 1910: 39 [40].Philolestes Kieffer, 1922: 205 [13].Psilogasteroides Brèthes, 1911: 209–210 [43].Distribution
Restricted to lowland rainforests of continental South America [5].Biology
Members ofNotoxoides display some of the most advanced associations with ants. So far, ants of genera Neivamyrmex and Eciton (Ecitonini) were recorded as hosts [19]. Adult wasps are frequently collected in light traps. Wings may be lost to typical alectomy as indicated by shriveled wing rudiments in some specimens [5].


### 3.1.9.Notoxoides pedissequus (Borgmeier, 1939) [44]

Notoxopria pedissequaBorgmeier, 1939: 538 [44].Distribution
Argentina (Córdoba) [19].Biology
Associated withNeivamyrmex pseudops (Forel) [44].Remarks
Loiácono [20] studied a female alate collected by Bruch in Córdoba province.


### 3.1.10.Notoxoides pronotalis(Borgmeier, 1939) [44] (Figures 4(a), 4(b), and 5)

Notoxoides pronotalis female in dorsal view: (a) alate and (b) dealate specimens. Scale: 1 mm. (c) Eciton dulciumcollected with Notoxoides pronotalis, in lateral view.Scale: 1 mm.
(a)
(b)
(c)Figure 5
Original illustration ofNotoxoides pronotalisfemale in lateral view, by Bruch.Philolestes rufus Kieffer, 1922: 205 [13].Philolestes pronotalis Borgmeier, 1939: 536 [44].Notoxoides pronotalis: Masner, 1977: 34 [45].Notoxoides kiefferi Loiácono, 1981: 305, 306 [19].Distribution
Argentina (Córdoba, Salta, San Luis, and Santiago del Estero) [19, 44].Biology
Associated with Eciton dulcium Forel andNeivamyrmex sulcatus (Mayr) [44].Material Studied
Syntype, female dealated, collected withEciton dulcium, Argentina, Córdoba, Alta Gracia, 4-XII-I921, Bruch coll.; 2 syntype females alated, same data as syntype except II-1922, collected with Neivamyrmex sulcatus, Bruch coll. and det.; 21 females dealated, Argentina, Salta, Tartagal, I-1960, Martínez coll., with a Eciton dulcium, and 5 females alated, Argentina, Salta, Pocitos, III-1959, Martínez coll.; 3 females dealated and 1 alated, Córdoba, San Javier, La Paz, 15-31-XII-1928, Bruch coll., with Eciton dulcium; Córdoba, Alta Gracia: 1 female dealated, collected with Eciton dulcium (Figure 4(c)), 4-XII-1922, Bruch coll.; 1 female dealated,  without date and collector; 1 female alated, La Granja, 21-VIII-1924, Bruch coll.; 2 females dealated, La Granja, 25-I-1925, Bruch coll.; 3 females alated, La Granja, 4-XI-1925, Bruch coll.; 2 females dealated, with Eciton dulcium, La Granja, 4-XI-1925, Bruch coll.; 2 females alated with Eciton dulcium, 13-III-1934, Bruch coll.;  3 females alated, without date, Bruch coll.; 1 female dealated; Córdoba, Unquillo, without date and collector; 9 females alated, Córdoba, Unquillo, without date and collector; 1 female with fore wings, Córdoba, Unquillo, without date and collector; 2 females alated, Santiago del Estero, Cerrillos, 2-V-1955, without collector, and 2 females alated, without date, Bruch coll.; 5 females alated, without locality, 21-II-1925, light collected, without collector; 2 females dealated and 3 alated, without locality, 22-II-1925, light collected, without collector; 5 females alated and 1 dealated, without locality, 23-II-1925, light collected, without collector; 1 female alated, without locality, 24-II-1925, light collected, without collector.Remarks
Bruch always sent to Kieffer diapriid samples to be studied. As we mentioned, he was an excellent scientific illustrator (Figure5) [46] and an important photographer as is shown in (Figure 6) Neivamyrmex pseudops, ant host of Notoxoides pedisequus [47].Figure 6
Neivamyrmex pseudops,gravid queen in dorsal view, photographed by Bruch.We observed numerous both alate and dealate individuals found dependent on the phase of life. As is mentioned [2], during the alate phase, numerous adults were caught by light traps as we observed in the female material light collected by Bruch.Lachaud [48] mentioned that ants search actively for some chemical substances produced by glands at the basis of the setae present on the diapriid cuticle; similarly we observed the presence of peculiar neck hairs in N. pronotalis [20].


### 3.1.11.Szelenyiopria Fabritius, 1974 [49]

SzelenyiopriaFabritius, 1974: 54 [49].Gymnopria Loiácono, 1987: 130 [21].Distribution
Wide distribution from Argentina to Guatemala [21, 49].Biology
Szelenyiopria lucens (Loiácono) from Uruguay is the first member of the tribe Diapriini in the New World positively reared from ants. Loiácono [21] reports up to three wasps per mature larva of Acromyrmex ambiguus (Emery) (Attini). Members of Szelenyiopria show no specialized structures known among other myrmecophilic Diapriini; Masner and García [5] assumed that the specialized setae with truncate apices are outlet of chemical substances.


### 3.1.12.Szelenyiopria pampeana (Loiácono, 2000) [4] (Figure 7(a))

(a)Szelenyiopria pampeanafemale in lateral view. (b) Acromyrmex lobicornis larva showing immature instars of diapriines.Scale: 1 mm.
(a)
(b)Gymnopria pampeana Loiácono, 2000: 10 in Loiácono et al., 2000 [4].Szelenyiopria pampeana: Loiácono and Margaría, 2009: 63 [8].Distribution
Argentina (La Pampa) [4, 8].Biology
Koinobiont and gregarious endoparasitoids of late instar larvae ofAcromyrmex lobicornis (Emery), it was also established simultaneous parasitoidism with Trichopria sp. [4] (Figure 7(b)).Material Studied
Holotype female, Argentina, Santa Rosa, 8-XI-1995, Quirán and Corró Molas colls.; 25 paratypes females and 3 males, Lihuel Calel, 4-XII-1997,  Quirán and Corró Molas colls.


### 3.1.13.Szelenyiopria reichenspergeri (Ferrière, 1929) [35]

Doliopria reinchespergeri Ferrière, 1929: 165 [35].Szelenyiopria reinchespergeri: Fabritius, 1974, 54 [49].Distribution
Argentina (Salta and Tucumán) [35, 49].Biology
Associated withEciton quadriglume and Neivamyrmex legionis (Smith) [35, 49].Material Studied
One female, Argentina, Salta, 2-6-II-1950, Golbach coll.


### 3.1.14.Szelenyiopria sp.

Distribution
Argentina (Córdoba) (new record).Material Studied
Female and 3 males with an ecitonine ant, Argentina, Córdoba, San Javier, La Paz, 1-20-I-1929, Bruch coll.Remarks
Most females of this genus have 11-segmented antennae, but material studied here presents antenna 12-segmented as mentioned by Masner and García [5] for undescribed species. We considered that these specimens belong to Szelenyiopria genus by the most important feature, the presence on entire body of specialized straight setae, truncate apically.


### 3.1.15.Trichopria Ashmead, 1893 [34]

Trichopria Ashmead, 1893: 407, 431 [34].Ashmeadopria Kieffer, 1912: 8, 10, 59 [50].Phaenopria Ashmead, 1893: 40, 436 [34].Planopria Kieffer, 1906: 19 [51].Orthopria Kieffer, 1911: 983, 984 [52].Distribution
Worldwide [5].Biology
Associated with the “fire ant,”Solenopsis richteri Forel (Kieffer, 1921) and endoparasitoid of Acromyrmex lobicornis [4].


### 3.1.16.Trichopria formicans Loiácono, 2000 [4] (Figures 8(a) and 8(b))

Trichopria formicans female (a) in dorsal view and (b) lateral view. Scale: 1 mm.
(a)
(b)Trichopria formicans  Loiácono 2000 in Loiácono et al., 2000: 12 [4].Distribution
Argentina (La Pampa) [4].Biology
Reared from larvae ofAcromyrmex lobicornis [4].Material Studied
Holotype female, Argentina, La Pampa, Utracán, 22-XII-1997, Caramuti y Rodriguez colls.; paratypes 68 females and 43 males (MLP), same data as holotype.


### 3.1.17.Trichopria myrmecophila (Kieffer, 1921) [12]

Phaenopria myrmecophila Kieffer, 1921: 4 [12].Trichopria myrmecophila: De Santis in De Santis and Esquivel, 1966: 50 [16].Distribution
Argentina (Buenos Aires) [12].Biology
Associated withSolenopsis richteri [12].


### 3.1.18.Trichopria sp.

Distribution
Argentina Buenos Aires.Biology
Collected with the “argentine ant,”Linepithema humile (new record).Material Studied
Female collected withLinepithema humile, Argentina, Buenos Aires, J. C. Paz, 8-II-1940, Ogloblin coll.Remarks
Masner studied this material and determined specimens asTrichopria s. str. sp.


## 3.1.1.Asolenopsia Kieffer, 1921 [12]

Asolenopsia Kieffer, 1921: 36 [12].Euplacopria Ferrière, 1929: 157 [35].Distribution
Tropical lowlands of Central and South America [5].Biology
Associated with ecitonini ants of genusEciton Latreille, Labidus Jurine  and Neivamyrmex Borgmeier [5].Remarks
Members ofAsolenopsia are moderately to highly specialized associates to ecitonine ants [20]. Their wings are primarily developed but subsequently bitten off by ants or cast off spontaneously (alectomy). Winged adults are also collected in light traps [5].


## 3.1.2.Asolenopsia rufa Kieffer, 1921 [12] (Figure 1(a))

(a)Asolenopsia rufa female dealate in dorsal view. (b) Neivamyrmex carettei.Scale: 1 mm.
(a)
(b)Asolenopsia rufa Kieffer, 1921: 37 [12].Distribution
Argentina (Córdoba, Entre Ríos, and Santa Fe) [12, 17].Biology
Associated withNeivamyrmex carettei (Forel) [12] (Figure 1(b)).Material Studied
Syntype, female, dealated, withNeivamyrmex carettei worker, Argentina, Córdoba, Alta Gracia, La Granja, 1-8-IV-1920, Bruch coll.; one female, without date, Santa Fe, Vera y Pintado (Fives Lille), Weiser coll.; female alated, Argentina, Misiones, Loreto, without date, Ogloblin coll.


## 3.1.3.Basalys Westwood, 1832 [36]

BasalysWestwood, 1832: 342–344 [36].CeratopriaAshmead, 1893: 407, 42 [34].Acidopria Kieffer, 1913: 442 [37].Loxotropa auct. nec Foerster, synonymized by Masner, 1964 [38].Nesopria Muesebeck and Walkley, 1956: 319–419 [39].Distribution
The genus is well represented in North and South America, rarely in Chile [5].Biology
Several species were reared from various dipterous hosts, and some were collected in ant nests [5].


## 3.1.4.Basalys sp.

Material Studied
One female and 1 male (microscopic slide) collected with the “Argentine ant,”Linepithema humile (Mayr), Argentina, Buenos Aires, J. C. Paz, 11-X-1934, Ogloblin coll.; 1 female (microscopic slide) collected with the Argentine ant Linepithema humile, Argentina, Buenos Aires, J. C. Paz, 8-IX-1945, Bezzi leg.Biology
Associated withLinepithema humile (new record).Remarks
Female and male studied were determined by Masner, who wrote a label: “Basalys sp. ♀♂(=Loxotropa auct.) aberrant sp. with !11-segm. ant. ♀, Det. L. Masner, ‘89”; and female specimen: “Basalys sp. ♀(=Loxotropa auct.) !11-segmented antenna, Det. L. Masner, ’89.” Specimens studied were determined by Ogloblin as a new species of Doliopria, but he did not describe it. We also considered that material studied belong to genus Basalys, as it was established by Masner.


## 3.1.5.Doliopria Kieffer, 1910 [40]

Doliopria Kieffer, 1910: 48 [40].Martinica Risbec, 1950: 533 [41].Distribution
Doliopria is restricted to the New World, with only a few species in the Nearctic region and with a high number of undescribed species in tropical America [5].Biology
Associated with ecitonini and attini ants [5].Remarks
Three Neotropical species were described associated with ants [12, 35, 40]; hypothetically they parasitized synoeketic Diptera because they show no specialized morphology [5].


## 3.1.6.Doliopria collegii Ferrière, 1929 [35] (Figure 2)

Figure 2
Doliopria collegii female in lateral view. Scale: 1 mm.Doliopria collegii Ferrière, 1929 : 164 [35].Distribution
Argentina (Buenos Aires and Misiones) [18, 35].Biology
Associated with ecitonini ants,Eciton burchellii (Westwood) and Eciton quadriglume (Haliday) [35].Material Studied
Two females alated, Argentina, Misiones, Loreto, 20-X-1919 and 18-IX-1923, Ogloblin coll. and det.


## 3.1.7.Doliopria myrmecobia Kieffer, 1921 [12] (Figure 3(a))

(a)Doliopria myrmecobia female in lateral view. (b) Acromyrmex lundii.Scale: 1 mm.
(a)
(b)Doliopria myrmecobia Kieffer, 1921: 39 [12].Distribution
Argentina (Buenos Aires; Misiones, new record) [12].Biology
Associated with attini antsAcromyrmex lundii (Guérin-Méneville) [12] (Figure 3(b)).Material Studied
One female, Argentina, Buenos Aires, La Plata, VIII, inside a nest ofAcromyrmex lundii, Bruch coll.; 1 female, alated collected with Acromyrmex sp., Argentina, Misiones, Loreto, 3-XI-1928, Ogloblin coll. and det.


## 3.1.8.Notoxoides Ashmead, 1903 [42]

Notoxoides Ashmead, 1903: 30 [42].NotoxopriaKieffer, 1910: 39 [40].Philolestes Kieffer, 1922: 205 [13].Psilogasteroides Brèthes, 1911: 209–210 [43].Distribution
Restricted to lowland rainforests of continental South America [5].Biology
Members ofNotoxoides display some of the most advanced associations with ants. So far, ants of genera Neivamyrmex and Eciton (Ecitonini) were recorded as hosts [19]. Adult wasps are frequently collected in light traps. Wings may be lost to typical alectomy as indicated by shriveled wing rudiments in some specimens [5].


## 3.1.9.Notoxoides pedissequus (Borgmeier, 1939) [44]

Notoxopria pedissequaBorgmeier, 1939: 538 [44].Distribution
Argentina (Córdoba) [19].Biology
Associated withNeivamyrmex pseudops (Forel) [44].Remarks
Loiácono [20] studied a female alate collected by Bruch in Córdoba province.


## 3.1.10.Notoxoides pronotalis(Borgmeier, 1939) [44] (Figures 4(a), 4(b), and 5)

Notoxoides pronotalis female in dorsal view: (a) alate and (b) dealate specimens. Scale: 1 mm. (c) Eciton dulciumcollected with Notoxoides pronotalis, in lateral view.Scale: 1 mm.
(a)
(b)
(c)Figure 5
Original illustration ofNotoxoides pronotalisfemale in lateral view, by Bruch.Philolestes rufus Kieffer, 1922: 205 [13].Philolestes pronotalis Borgmeier, 1939: 536 [44].Notoxoides pronotalis: Masner, 1977: 34 [45].Notoxoides kiefferi Loiácono, 1981: 305, 306 [19].Distribution
Argentina (Córdoba, Salta, San Luis, and Santiago del Estero) [19, 44].Biology
Associated with Eciton dulcium Forel andNeivamyrmex sulcatus (Mayr) [44].Material Studied
Syntype, female dealated, collected withEciton dulcium, Argentina, Córdoba, Alta Gracia, 4-XII-I921, Bruch coll.; 2 syntype females alated, same data as syntype except II-1922, collected with Neivamyrmex sulcatus, Bruch coll. and det.; 21 females dealated, Argentina, Salta, Tartagal, I-1960, Martínez coll., with a Eciton dulcium, and 5 females alated, Argentina, Salta, Pocitos, III-1959, Martínez coll.; 3 females dealated and 1 alated, Córdoba, San Javier, La Paz, 15-31-XII-1928, Bruch coll., with Eciton dulcium; Córdoba, Alta Gracia: 1 female dealated, collected with Eciton dulcium (Figure 4(c)), 4-XII-1922, Bruch coll.; 1 female dealated,  without date and collector; 1 female alated, La Granja, 21-VIII-1924, Bruch coll.; 2 females dealated, La Granja, 25-I-1925, Bruch coll.; 3 females alated, La Granja, 4-XI-1925, Bruch coll.; 2 females dealated, with Eciton dulcium, La Granja, 4-XI-1925, Bruch coll.; 2 females alated with Eciton dulcium, 13-III-1934, Bruch coll.;  3 females alated, without date, Bruch coll.; 1 female dealated; Córdoba, Unquillo, without date and collector; 9 females alated, Córdoba, Unquillo, without date and collector; 1 female with fore wings, Córdoba, Unquillo, without date and collector; 2 females alated, Santiago del Estero, Cerrillos, 2-V-1955, without collector, and 2 females alated, without date, Bruch coll.; 5 females alated, without locality, 21-II-1925, light collected, without collector; 2 females dealated and 3 alated, without locality, 22-II-1925, light collected, without collector; 5 females alated and 1 dealated, without locality, 23-II-1925, light collected, without collector; 1 female alated, without locality, 24-II-1925, light collected, without collector.Remarks
Bruch always sent to Kieffer diapriid samples to be studied. As we mentioned, he was an excellent scientific illustrator (Figure5) [46] and an important photographer as is shown in (Figure 6) Neivamyrmex pseudops, ant host of Notoxoides pedisequus [47].Figure 6
Neivamyrmex pseudops,gravid queen in dorsal view, photographed by Bruch.We observed numerous both alate and dealate individuals found dependent on the phase of life. As is mentioned [2], during the alate phase, numerous adults were caught by light traps as we observed in the female material light collected by Bruch.Lachaud [48] mentioned that ants search actively for some chemical substances produced by glands at the basis of the setae present on the diapriid cuticle; similarly we observed the presence of peculiar neck hairs in N. pronotalis [20].


## 3.1.11.Szelenyiopria Fabritius, 1974 [49]

SzelenyiopriaFabritius, 1974: 54 [49].Gymnopria Loiácono, 1987: 130 [21].Distribution
Wide distribution from Argentina to Guatemala [21, 49].Biology
Szelenyiopria lucens (Loiácono) from Uruguay is the first member of the tribe Diapriini in the New World positively reared from ants. Loiácono [21] reports up to three wasps per mature larva of Acromyrmex ambiguus (Emery) (Attini). Members of Szelenyiopria show no specialized structures known among other myrmecophilic Diapriini; Masner and García [5] assumed that the specialized setae with truncate apices are outlet of chemical substances.


## 3.1.12.Szelenyiopria pampeana (Loiácono, 2000) [4] (Figure 7(a))

(a)Szelenyiopria pampeanafemale in lateral view. (b) Acromyrmex lobicornis larva showing immature instars of diapriines.Scale: 1 mm.
(a)
(b)Gymnopria pampeana Loiácono, 2000: 10 in Loiácono et al., 2000 [4].Szelenyiopria pampeana: Loiácono and Margaría, 2009: 63 [8].Distribution
Argentina (La Pampa) [4, 8].Biology
Koinobiont and gregarious endoparasitoids of late instar larvae ofAcromyrmex lobicornis (Emery), it was also established simultaneous parasitoidism with Trichopria sp. [4] (Figure 7(b)).Material Studied
Holotype female, Argentina, Santa Rosa, 8-XI-1995, Quirán and Corró Molas colls.; 25 paratypes females and 3 males, Lihuel Calel, 4-XII-1997,  Quirán and Corró Molas colls.


## 3.1.13.Szelenyiopria reichenspergeri (Ferrière, 1929) [35]

Doliopria reinchespergeri Ferrière, 1929: 165 [35].Szelenyiopria reinchespergeri: Fabritius, 1974, 54 [49].Distribution
Argentina (Salta and Tucumán) [35, 49].Biology
Associated withEciton quadriglume and Neivamyrmex legionis (Smith) [35, 49].Material Studied
One female, Argentina, Salta, 2-6-II-1950, Golbach coll.


## 3.1.14.Szelenyiopria sp.

Distribution
Argentina (Córdoba) (new record).Material Studied
Female and 3 males with an ecitonine ant, Argentina, Córdoba, San Javier, La Paz, 1-20-I-1929, Bruch coll.Remarks
Most females of this genus have 11-segmented antennae, but material studied here presents antenna 12-segmented as mentioned by Masner and García [5] for undescribed species. We considered that these specimens belong to Szelenyiopria genus by the most important feature, the presence on entire body of specialized straight setae, truncate apically.


## 3.1.15.Trichopria Ashmead, 1893 [34]

Trichopria Ashmead, 1893: 407, 431 [34].Ashmeadopria Kieffer, 1912: 8, 10, 59 [50].Phaenopria Ashmead, 1893: 40, 436 [34].Planopria Kieffer, 1906: 19 [51].Orthopria Kieffer, 1911: 983, 984 [52].Distribution
Worldwide [5].Biology
Associated with the “fire ant,”Solenopsis richteri Forel (Kieffer, 1921) and endoparasitoid of Acromyrmex lobicornis [4].


## 3.1.16.Trichopria formicans Loiácono, 2000 [4] (Figures 8(a) and 8(b))

Trichopria formicans female (a) in dorsal view and (b) lateral view. Scale: 1 mm.
(a)
(b)Trichopria formicans  Loiácono 2000 in Loiácono et al., 2000: 12 [4].Distribution
Argentina (La Pampa) [4].Biology
Reared from larvae ofAcromyrmex lobicornis [4].Material Studied
Holotype female, Argentina, La Pampa, Utracán, 22-XII-1997, Caramuti y Rodriguez colls.; paratypes 68 females and 43 males (MLP), same data as holotype.


## 3.1.17.Trichopria myrmecophila (Kieffer, 1921) [12]

Phaenopria myrmecophila Kieffer, 1921: 4 [12].Trichopria myrmecophila: De Santis in De Santis and Esquivel, 1966: 50 [16].Distribution
Argentina (Buenos Aires) [12].Biology
Associated withSolenopsis richteri [12].


## 3.1.18.Trichopria sp.

Distribution
Argentina Buenos Aires.Biology
Collected with the “argentine ant,”Linepithema humile (new record).Material Studied
Female collected withLinepithema humile, Argentina, Buenos Aires, J. C. Paz, 8-II-1940, Ogloblin coll.Remarks
Masner studied this material and determined specimens asTrichopria s. str. sp.


## 3.2. Tribe Spilomicrini Ashmead, 1893 [34]

### 3.2.1.BruchopriaKieffer, 1921 [12]

Bruchopria Kieffer, 1921: 38 [12].Aulatopria Brèthes, 1927: 164 [53].Distribution
Argentina (Buenos Aires, Córdoba, and Misiones) [12, 53].Biology
Associated with ants of the generaSolenopsis Westwood (Solenopsidini) and Acromyrmex  Mayr (Attini) [12].Remarks
Hölldobler and Wilson [54] mentioned specimens of genus Bruchopria, as Solenopsis guest. Masner and García [5] mentioned “wings often bitten off by ants.” Loiácono et al. [26] studied alated and dealated individuals of Bruchopria species. The action of dealation has not been observed. The presence of tegulae with normal development and wing stumps demonstrates that the apterism has a secondary origin, caused by the autotomy or by bites of the host ants. The apices of the wing stumps of all individuals examined were regular suggesting that the wings are bitten or torn off close to the tegulae. The fact that specimens are dealated allows them to move into the mound galleries and chambers.


### 3.2.2.Bruchopria hexatoma Kieffer, 1921 [12] (Figures 9(a), 10(a), and 10(b))

(a)Bruchopria hexatoma female dealate in dorsal view. (b) Solenopsis richteri.Scale: 1 mm.
(a)
(b)Bruchopria hexatoma female. (a) mesosoma and petiole in dorsal view, scale: 0.5 mm; (b) wing stump, scale: 0.1 mm [26].
(a)
(b)Bruchopria hexatoma
Kieffer, 1921: 39 [12].Bruchopria hexatoma: Borgmeier, 1939: 543 [44].Distribution
Argentina (Misiones, Córdoba and Buenos Aires) [12, 44].Biology
Associated withSolenopsis richteri (Figure 9(b)) and Acromyrmex lundii [12, 44].Material Studied
One female dealated, Argentina, Misiones, Pastoreo Grande, 9-VII-1932, Ogloblin coll.; 1 female dealated, Argentina, Córdoba, XII-1920, Bruch coll., 1 female dealated, Córdoba, Sierras de Córdoba, La Granja, Bruch coll., without date; 1 male dealated, Argentina, Buenos Aires, without locality, 9-VII-1923, Bruch coll., with the ant; 4 females dealated, Argentina, Buenos Aires, Olivos, without date, Bruch coll., with the ant; 1 female dealated, Argentina, Buenos Aires, 10-IX-1925, Bruch coll.; 1 female dealated withAcromyrmex lundii, Argentina, Buenos Aires, without date, Bruch coll.Remarks
Bruchopria hexatoma has been reported by Kieffer [12] in association with Solenopsis richteri and Acromyrmex lundii in Argentina; Borgmeier [44] also mentioned this species as a guest of S. saevissima (Smith), in Brazil.The specimens from the provinces of Córdoba and Buenos Aires are dealated, with remains of wings (Figures10(a) and 10(b)), and most of them are accompanied by the host ants (Figure 9(b)). Unfortunately, the types of the species described by Kieffer have become widely scattered or lost [55]. Bruch sent to Kieffer part of the same series of material to identify (De Santis, pers. comm.).


### 3.2.3.Bruchopria pentatoma Kieffer, 1921 [12]

Bruchopria pentatoma Kieffer, 1921: 38 [12].Distribution
Argentina (Córdoba) [12].Biology
Associated withSolenopsis richteri [12].Material Studied
Syntype male dealated, Argentina, Córdoba, Alta Gracia; 1-8-IV-1920, Bruch coll.Remarks
According to Kieffer's description, females of both species,B. pentatoma and B. hexatoma, are distinguished by the number of club antennomeres, five and six, respectively. Unfortunately, the unique female type is not available. Bruchopria pentatoma has also been reported by Kieffer [12] in association with S. richteri and Acromyrmex lundii (Guérin) in Argentina.


### 3.2.4.PentapriaKieffer, 1905 [56]

Pentapria Kieffer, 1905: 34 [56].Antipapria Fabritius, 1968: 844 [57].Bakeria Kieffer, 1905: 34 [56].Plutopria Kieffer, 1910: 48 [40].Spilomicrinus Ogloblin, 1957: 425 [58].Xenopria Fouts, 1939: 260 [59].Distribution
The genus is distributed in the New World [5].Biology
The principal host plausible to assume is Stratiomyidae (Diptera) [5]. Herein, we studied a female collected with Solenopsis saevissima (Hymenoptera: Formicidae).


### 3.2.5.Pentapria cf. nodicornis

Distribution
Argentina (Córdoba).Biology
Associated withSolenopsis saevissima (new record).Material Studied
Female collected withSolenopsis saevissima, Argentina, Córdoba, Alta Gracia, La Granja, II-1927, Bruch. coll., with no more data.


### 3.2.6.SpilomicrusWestwood, 1832 [36]

SpilomicrusWestwood, 1832: 129 [36].LoxotropaFoerster, 1856: 122, 123, 126 [60].HoplopriaAshmead, 1893: 385, 386, 388 [34].LinkiolaKieffer, 1910: 39 [40].EriopriaKieffer, 1910: 693, 744 [40].TritopriaKieffer, 1910: 717, 748 [40].CologlyptusCrawford, 1910: 123 [61].ScutellipriaSzabó, 1961: 53–493 [62].Distribution
America [5].Biology
Primary parasitoidism solitary and gregarious of various Diptera; few species were reared from Coleoptera [5]. Herein, we studied samples associated with a Solenopsidini ant.


### 3.2.7.Spilomicrus sp.

Distribution
Argentina (Buenos Aires).Biology
Associated with Solenopsidini ant.Material Studied
Two females with a Solenopsidini ant, Argentina, Buenos Aires, 9-VIII-1923, Bruch coll.Table1 summarizes information about diapriids and their associates.Table 1
Diapriid tribe
Diapriids species
Argentine provinces
Ant subfamily
Ant tribe
Ant species
Diapriini
Asolenopsia rufa
Córdoba, Entre Ríos, Santa Fe
Ecitoninae
Ecitonini
Neivamyrmex carettei
Basalyssp.
Buenos Aires
Dolichoderinae
Dolichoderini
Linepithema humile
Doliopria collegii
Buenos Aires, Misiones
Ecitoninae
Ecitonini
Eciton burchellii, Eciton quadriglume
Doliopria myrmecobia
Buenos Aires, Misiones
Myrmicinae
Attini
Acromyrmex lundii
Notoxoides pedissequus
Córdoba
Ecitoninae
Ecitonini
Neivamyrmex pseudops
Notoxoides pronotalis
Córdoba, Salta, San Luis,  Santiago del Estero
Ecitoninae
Ecitonini
Eciton dulcium, Neivamyrmex sulcatus
Szelenyiopria pampeana
La Pampa
Myrmicinae
Attini
Acromyrmex lobicornis
Szelenyiopria reichenspergeri
Salta, Tucumán
Ecitoninae
Ecitonini
Eciton quadriglume, Neivamyrmex legionis
Szelenyiopriasp.
Córdoba
Ecitoninae
Ecitonini
Ecitonini sp.
Trichopria formicans
La Pampa
Myrmicinae
Attini
Acromyrmex lobicornis
Trichopria myrmecophila
Buenos Aires
Myrmicinae
Solenopsidini
Solenopsis richteri
Trichopria sp.
Buenos Aires
Dolichoderinae
Dolichoderini
Linepithema humile
Spilomicrini
Bruchopria hexatoma
Buenos Aires, Córdoba, Misiones
MyrmicinaeMyrmicinae
SolenopsidiniAttini
Solenopsis richteri
Acromyrmex lundii
Bruchopria pentatoma
Córdoba
Myrmicinae
Solenopsidini
Solenopsis richteri
Pentapria cf. nodicornis
Córdoba
Myrmicinae
Solenopsidini
Solenopsis saevissima
Spilomicrussp.
Buenos Aires
Myrmicinae
Solenopsidini
Solenopsidini sp.


## 3.2.1.BruchopriaKieffer, 1921 [12]

Bruchopria Kieffer, 1921: 38 [12].Aulatopria Brèthes, 1927: 164 [53].Distribution
Argentina (Buenos Aires, Córdoba, and Misiones) [12, 53].Biology
Associated with ants of the generaSolenopsis Westwood (Solenopsidini) and Acromyrmex  Mayr (Attini) [12].Remarks
Hölldobler and Wilson [54] mentioned specimens of genus Bruchopria, as Solenopsis guest. Masner and García [5] mentioned “wings often bitten off by ants.” Loiácono et al. [26] studied alated and dealated individuals of Bruchopria species. The action of dealation has not been observed. The presence of tegulae with normal development and wing stumps demonstrates that the apterism has a secondary origin, caused by the autotomy or by bites of the host ants. The apices of the wing stumps of all individuals examined were regular suggesting that the wings are bitten or torn off close to the tegulae. The fact that specimens are dealated allows them to move into the mound galleries and chambers.


## 3.2.2.Bruchopria hexatoma Kieffer, 1921 [12] (Figures 9(a), 10(a), and 10(b))

(a)Bruchopria hexatoma female dealate in dorsal view. (b) Solenopsis richteri.Scale: 1 mm.
(a)
(b)Bruchopria hexatoma female. (a) mesosoma and petiole in dorsal view, scale: 0.5 mm; (b) wing stump, scale: 0.1 mm [26].
(a)
(b)Bruchopria hexatoma
Kieffer, 1921: 39 [12].Bruchopria hexatoma: Borgmeier, 1939: 543 [44].Distribution
Argentina (Misiones, Córdoba and Buenos Aires) [12, 44].Biology
Associated withSolenopsis richteri (Figure 9(b)) and Acromyrmex lundii [12, 44].Material Studied
One female dealated, Argentina, Misiones, Pastoreo Grande, 9-VII-1932, Ogloblin coll.; 1 female dealated, Argentina, Córdoba, XII-1920, Bruch coll., 1 female dealated, Córdoba, Sierras de Córdoba, La Granja, Bruch coll., without date; 1 male dealated, Argentina, Buenos Aires, without locality, 9-VII-1923, Bruch coll., with the ant; 4 females dealated, Argentina, Buenos Aires, Olivos, without date, Bruch coll., with the ant; 1 female dealated, Argentina, Buenos Aires, 10-IX-1925, Bruch coll.; 1 female dealated withAcromyrmex lundii, Argentina, Buenos Aires, without date, Bruch coll.Remarks
Bruchopria hexatoma has been reported by Kieffer [12] in association with Solenopsis richteri and Acromyrmex lundii in Argentina; Borgmeier [44] also mentioned this species as a guest of S. saevissima (Smith), in Brazil.The specimens from the provinces of Córdoba and Buenos Aires are dealated, with remains of wings (Figures10(a) and 10(b)), and most of them are accompanied by the host ants (Figure 9(b)). Unfortunately, the types of the species described by Kieffer have become widely scattered or lost [55]. Bruch sent to Kieffer part of the same series of material to identify (De Santis, pers. comm.).


## 3.2.3.Bruchopria pentatoma Kieffer, 1921 [12]

Bruchopria pentatoma Kieffer, 1921: 38 [12].Distribution
Argentina (Córdoba) [12].Biology
Associated withSolenopsis richteri [12].Material Studied
Syntype male dealated, Argentina, Córdoba, Alta Gracia; 1-8-IV-1920, Bruch coll.Remarks
According to Kieffer's description, females of both species,B. pentatoma and B. hexatoma, are distinguished by the number of club antennomeres, five and six, respectively. Unfortunately, the unique female type is not available. Bruchopria pentatoma has also been reported by Kieffer [12] in association with S. richteri and Acromyrmex lundii (Guérin) in Argentina.


## 3.2.4.PentapriaKieffer, 1905 [56]

Pentapria Kieffer, 1905: 34 [56].Antipapria Fabritius, 1968: 844 [57].Bakeria Kieffer, 1905: 34 [56].Plutopria Kieffer, 1910: 48 [40].Spilomicrinus Ogloblin, 1957: 425 [58].Xenopria Fouts, 1939: 260 [59].Distribution
The genus is distributed in the New World [5].Biology
The principal host plausible to assume is Stratiomyidae (Diptera) [5]. Herein, we studied a female collected with Solenopsis saevissima (Hymenoptera: Formicidae).


## 3.2.5.Pentapria cf. nodicornis

Distribution
Argentina (Córdoba).Biology
Associated withSolenopsis saevissima (new record).Material Studied
Female collected withSolenopsis saevissima, Argentina, Córdoba, Alta Gracia, La Granja, II-1927, Bruch. coll., with no more data.


## 3.2.6.SpilomicrusWestwood, 1832 [36]

SpilomicrusWestwood, 1832: 129 [36].LoxotropaFoerster, 1856: 122, 123, 126 [60].HoplopriaAshmead, 1893: 385, 386, 388 [34].LinkiolaKieffer, 1910: 39 [40].EriopriaKieffer, 1910: 693, 744 [40].TritopriaKieffer, 1910: 717, 748 [40].CologlyptusCrawford, 1910: 123 [61].ScutellipriaSzabó, 1961: 53–493 [62].Distribution
America [5].Biology
Primary parasitoidism solitary and gregarious of various Diptera; few species were reared from Coleoptera [5]. Herein, we studied samples associated with a Solenopsidini ant.


## 3.2.7.Spilomicrus sp.

Distribution
Argentina (Buenos Aires).Biology
Associated with Solenopsidini ant.Material Studied
Two females with a Solenopsidini ant, Argentina, Buenos Aires, 9-VIII-1923, Bruch coll.Table1 summarizes information about diapriids and their associates.Table 1
Diapriid tribe
Diapriids species
Argentine provinces
Ant subfamily
Ant tribe
Ant species
Diapriini
Asolenopsia rufa
Córdoba, Entre Ríos, Santa Fe
Ecitoninae
Ecitonini
Neivamyrmex carettei
Basalyssp.
Buenos Aires
Dolichoderinae
Dolichoderini
Linepithema humile
Doliopria collegii
Buenos Aires, Misiones
Ecitoninae
Ecitonini
Eciton burchellii, Eciton quadriglume
Doliopria myrmecobia
Buenos Aires, Misiones
Myrmicinae
Attini
Acromyrmex lundii
Notoxoides pedissequus
Córdoba
Ecitoninae
Ecitonini
Neivamyrmex pseudops
Notoxoides pronotalis
Córdoba, Salta, San Luis,  Santiago del Estero
Ecitoninae
Ecitonini
Eciton dulcium, Neivamyrmex sulcatus
Szelenyiopria pampeana
La Pampa
Myrmicinae
Attini
Acromyrmex lobicornis
Szelenyiopria reichenspergeri
Salta, Tucumán
Ecitoninae
Ecitonini
Eciton quadriglume, Neivamyrmex legionis
Szelenyiopriasp.
Córdoba
Ecitoninae
Ecitonini
Ecitonini sp.
Trichopria formicans
La Pampa
Myrmicinae
Attini
Acromyrmex lobicornis
Trichopria myrmecophila
Buenos Aires
Myrmicinae
Solenopsidini
Solenopsis richteri
Trichopria sp.
Buenos Aires
Dolichoderinae
Dolichoderini
Linepithema humile
Spilomicrini
Bruchopria hexatoma
Buenos Aires, Córdoba, Misiones
MyrmicinaeMyrmicinae
SolenopsidiniAttini
Solenopsis richteri
Acromyrmex lundii
Bruchopria pentatoma
Córdoba
Myrmicinae
Solenopsidini
Solenopsis richteri
Pentapria cf. nodicornis
Córdoba
Myrmicinae
Solenopsidini
Solenopsis saevissima
Spilomicrussp.
Buenos Aires
Myrmicinae
Solenopsidini
Solenopsidini sp.


## 4. Discussion

The knowledge of the biology and behavior of these myrmecophilic diapriids and the nature of their interactions with ants has progressed in Argentina since 1980 [63] to present. There are nine genera recorded from Argentina, which represents about 50% of the genera mentioned by Masner and García [5] from the New World.The study of Diapriidae Collection housed at División Entomología of Museo de La Plata, which includes Bruch and Ogloblin myrmecophilic diapriid specimens, allowed us to report 16 species of nine genera of Diapriinae associated with ants in Argentina. It is interesting to highlight thatAsolenopsia rufa, Notoxoides pronotalis, Bruchopria pentatoma,andB. hexatoma are the species with a high degree of integration with the host ants, adaptation being both morphological and behavioral.We mentioned for the first time the associations between the “argentine ant,”Linepithema humile,and both Basalys sp. and Trichopriasp., Pentapriacf.nodicornisand Solenopsis saevissima, and Spilomicrussp. and Solenopsidini ant.Doliopria myrmecobiais a new record  to Misiones. The only described species of Szelenyiopriaoccurs in La Pampa province, S. pampeana; an undescribed species is known to us from Córdoba.We considered thatSzelenyiopria pampeanaand Trichopria formicansparasitoids of Acromyrmex species in Argentina seem excellent potential models to explore how parasitoids impact ant colony demography, population biology, and ant community structure.



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*Source: 320590-2013-03-24.xml*