Zootaxa 3878 (1): 001–018
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Copyright © 2014 Magnolia Press
ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3878.1.1
http://zoobank.org/urn:lsid:zoobank.org:pub:FFAA0DB9-C6A4-4963-B60A-2BA9069951A8
Review of the genus Eotilla Schuster, 1949 (Hymenoptera: Bradynobaenidae:
Typhoctinae: Eotillini) and description of new species from Argentina
JAVIER TORRÉNS1,4, PATRICIO FIDALGO1, ARTURO ROIG-ALSINA2 & DENIS J. BROTHERS3
1
CRILAR-CONICET, Entre Ríos y Mendoza, 5301 Anillaco, La Rioja, Argentina.
E-mail: jtorrens@crilar-conicet.gob.ar; pfidalgo@crilar-conicet.gob.ar
2
Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Av. Angel Gallardo 470, 1405 Buenos Aires, Argentina.
E-mail: arturo@macn.gov.ar
3
School of Life Sciences, University of KwaZulu-Natal, Pietermaritzburg, Private Bag X01, Scottsville, 3209 South Africa.
E-mail: brothers@ukzn.ac.za
4
Corresponding author
Abstract
Diagnoses for the genus Eotilla Schuster, 1949 based on males and females, new generic and specific characters, descriptions of two new Argentinian species (Eotilla schusteri Torréns, Fidalgo, Roig-Alsina & Brothers, sp. nov. and E. medanito Torréns, Fidalgo, Roig-Alsina & Brothers, sp. nov., based on both sexes), a description of the female of E. superba
Brothers, 1974 and redescriptions of the males of E. mickeli Schuster, 1949 and E. superba are provided. A key to the
species of Eotilla is presented.
Key words: Eotilla, diagnosis, key to species
Introduction
Schuster (1949) described Eotilla and Prototilla and placed them in a new subfamily (Eotillinae) within the family
Mutillidae, along with four other subfamilies (Typhoctinae, Apterogyninae, Sphaeropthalminae and Mutillinae)
found in the Neotropical region. In phylogenetic studies of Hymenoptera Aculeata, Brothers (1975, 1999) and
Brothers and Carpenter (1993) placed Eotillini and Typhoctini as tribes of Typhoctinae, but distant from Mutillidae
and relocated within Bradynobaenidae (with Chyphotinae, Apterogyninae and Bradynobaeninae).
Recent molecular phylogenetic studies of the superfamily Vespoidea by Pilgrim et al. (2008) indicate that the
family Bradynobaenidae—according to the concept of Brothers (1975)—is paraphyletic; they placed
Apterogyninae and Bradynobaeninae in Bradynobaenidae within a superfamily Scolioidea, and Chyphotinae and
Typhoctinae in Chyphotidae within a superfamily Thynnoidea. This result was utilized by Debevec et al. (2012).
Heraty et al. (2011) in their study of the phylogeny of Hymenoptera, confirmed the lack of a close relationship
between Mutillidae and Chyphotes Blake, 1886, but showed a close relationship between Chyphotes and Colocistis
Krombein, 1942 (their only representative of Tiphiidae). Since the positions of these groups remain in discussion,
however, we retain the generally used classification.
At present, the Typhoctinae contains two tribes: Typhoctini with Typhoctes Ashmead, 1899 (North and Central
America) and Typhoctoides Brothers, 1974 (Chile and Argentina) and Eotillini with Eotilla (Chile and Argentina)
and Prototilla (Argentina). The revision of Prototilla by Torréns et al. (2012) included a diagnosis of Eotillini and
a comparative table between the two genera of the tribe; two new species based on males and two new species
based on females were described and P. typhoctoides (Martinez & Fritz, 1974) (ex Eotilla typhoctoides) was
proposed as a new combination. The authors did not include sex associations because of extreme sexual
dimorphism, and collection data did not indicate particular associations.
Currently, Eotilla comprises two species: the type species, E. mickeli Schuster, 1949 (Chile) and E. superba
Brothers, 1974 (Argentina), both based on males. Mickel (1968), however, assigned a female from Argentina to E.
Accepted by A. Lelej: 6 Oct. 2014; published: 22 Oct. 2014
1
mickeli. Since then, several Argentinean specimens of both sexes of different species have been collected and we
take this opportunity to record these here.
In this paper, diagnoses of Eotilla based on males and females are presented, new generic and specific
characters are added, and two new species of Eotilla are described from Argentina based on both sexes: E.
schusteri Torréns, Fidalgo, Roig-Alsina & Brothers, sp. nov. and E. medanito Torréns, Fidalgo, Roig-Alsina &
Brothers, sp. nov.. A description of the female of E. superba and redescriptions of the males of E. mickeli and E.
superba are provided. A key to the species concerned is given.
Material and methods
JT and PF collected males and females using yellow pan traps containing water and a few drops of detergent buried
to soil level and cleared daily; in addition, females were collected with pitfall traps containing propylene glycol and
covered to prevent the entry of sand and larger animals. The collection methods for other specimens are mostly
unrecorded, although a few males were collected in Malaise traps (MT).
Most images were obtained using Leica QWin software operating on a Leica M420 Macroscope linked to a
Sony PowerHad DXC-950P digital video camera; and Leica Application Suite (version 3.5.0) software operating
on a Leica MZ12 stereomicroscope linked to a Leica DFC295 digital video camera. Images were enhanced with
COREL PHOTOPAINT and COREL DRAW (version 15); and some images were processed with the computer
program DEEP FOCUS by Stuart Ball. A few images were made using a Canon PowerShot G10 camera adapted to
a Wild M400 photomacroscope using a Clearshot 600 adapter kit (Alexis Scientific) and stacked using CombineZP
software (Hadley 2010). The distribution map was made using Simple Mappr (Shorthouse, 2010).
Specimens studied are deposited in the Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”,
Buenos Aires, Argentina (MACN); Instituto Fundación Miguel Lillo, Tucumán, Argentina (IFML); Instituto
Argentino de Investigación de Zonas Aridas, Mendoza, Argentina (IADIZA); Museo Nacional de Historia Natural,
Santiago, Chile (MNHNS); Los Angeles County Museum of Natural History, Los Angeles, California, USA
(LACM); California Academy of Sciences, San Francisco, California, USA (CAS); Canadian National Collection
of Insects, Ottawa, Ontario, Canada (CNC); Museum National d’Histoire Naturelle, Paris, France (MNHN);
Museum für Naturkunde, Berlin, and the collection of Denis J. Brothers, Pietermaritzburg, South Africa (DJBC)
which will be transferred to the Iziko South African Museum, Cape Town, South Africa (SAMC) in due course.
Morphological terms are from Brothers (1975), Reid (1941) and Torréns et al. (2012), with details on sculpture
from Eady (1968) and Harris (1979). The abbreviations F, S and T refer to numbered flagellomeres, metasomal
sterna and metasomal terga, respectively; LOL = lateral ocellar line, OOL = ocellocular line and POL = postocellar
line. Mesosomal length was measured between its extremities (excluding the anterior pronotal collar) in dorsal
view. Metasomal length was measured from the anterior extremity of the dorsal surface of T1 to the apex of T7, in
dorsal view, but adjusted to take into account telescoping of the sclerites so as to reflect the length in a natural
condition. Length of T1 was measured laterally, from the base to the posterodorsal apical extremity.
Eotilla Schuster, 1949
Eotilla Schuster, 1949: 75–80. Additional citations: Mickel (1968; description of female); Brothers (1970; mention); Brothers
(1974; description of male); Brothers (1975; subsequent description); Martinez & Fritz (1974, description of female);
Fernández (2001, 2002; list); Brothers (2006; key); Pilgrim et al. (2008; mention); Torréns & Roig-Alsina (2009;
mention); Torréns et al. (2012; diagnosis).
Type species: Eotilla mickeli Schuster, 1949, by original designation.
Diagnosis. Modified from Torréns et al. (2012).
Male. Macropterous. Head with ocelli. Antennal scape with a ventral longitudinal carina. Horizontal face of
prosternum weakly projecting anteriorly, without a projection posteriorly. Mid coxae moderately separated by a
distance equal to half maximum width of femur. Fore wing with two submarginal cells. Hind wing with median cell
obliquely truncate at apex, not reaching level of origin of distal hamuli-group; median cell with two veins arising
from it (RS and Cu). Tarsal claws smooth, without ventral tooth (see Fig. 11 in Schuster 1949). Genitalia with basal
ring anteroventrally broadly emarginate.
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Female. Apterous. Head without ocelli. Mandible with small preapical denticle on inner edge. Pronotum
trapezoidal, with sides strongly convergent to posterior margin; humeral tubercle scarcely developed. Mesopleural
longitudinal carina reaching to anterior half of sclerite or beyond but never complete. Mid coxae widely separated
by a distance of 1.3 × maximum width of mid femur. Tarsal claws smooth, without ventral tooth. S6 with lateroposterior margin with distinct denticles, although sometimes weak.
Key to species of Eotilla
Males
1.
2.
3.
-
Head and mesosoma dark brown to black; metasoma much paler (Fig. 3) . . . . . . . . . . . . . . . . . . . . . . . . . E. medanito sp. nov.
Head, mesosoma and metasoma dark brown to black (Figs 1, 2, 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
T1 and T2 with abundant scale-like setae on almost all surfaces in addition to apical fringe (Fig. 15). Frons with scattered erect
pale subplumose setae (Fig. 19) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. schusteri sp. nov.
T1 and T2 with few or without scale-like setae, except for apical fringe (Figs 9, 11). Frons with scattered erect dark subplumose setae (Figs 17, 18) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
T7 with an inverted-V-shaped depression medially. T2 with a group of scale-like setae on central area (Figs 6, 11). Posteriormost terga and sterna without short and coarse dark setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. superba Brothers
T7 flattish to convex medially, without an inverted-V-shaped depression. T2 without a group of scale-like setae on central area
(Figs 5, 9). Posteriormost terga and sterna with short and coarse dark setae (Figs 9, 10) . . . . . . . . . . . . . . E. mickeli Schuster
Females (unknown in E. mickeli)
1.
-
2.
-
Entire metasoma black or T1 and T2 brownish to reddish and remainder black (Figs 30, 31). T2 with abundant scale-like setae,
except on central area (Figs 26, 27, 30, 31) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. schusteri sp. nov.
Metasoma almost entirely light brown to yellowish or reddish, except sometimes dark brown posteriorly and/or anteriorly
(Figs 22–25, 28, 29). T2 at most with a group of scale-like setae on each side of posterior margin (Fig. 28) or without scale-like
setae (Figs 25, 29) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
T2 with a group of scale-like setae on each side of posterior margin (Figs 22, 28). General body color light brown to yellowish
(Figs 22, 28). Head with scale-like setae, except on gena (Fig. 33) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. superba Brothers
T2 entirely without scale-like setae (Figs 23–25, 29). General body color light brown to reddish (Figs 23–25, 29), but sometimes pleura, propodeum, legs and last two or three segments of metasoma with darker brown areas. Head with dense scalelike setae, including on gena (Figs 34, 35) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .E. medanito sp. nov.
Eotilla mickeli Schuster, 1949
(Figs 1, 5, 9, 10, 17)
Eotilla mickeli Schuster, 1949: 77–80; holotype ♂, Chile: Valparaíso, Olmué [32°59'45"S 71°11'29"W] (Museum of
Comparative Zoology, Cambridge, Massachusetts, USA), not examined.
Diagnosis. Male. Head, mesosoma and metasoma dark brown to black (Figs 1, 5). T1, T2 and S2 with a few scalelike setae only on posterior margin or without (Figs 9, 10). T7 convex, without any inverted-V-shaped depression.
Posteriormost terga and sterna with dark, short and coarse setae (Figs 9, 10). Female. Unknown.
Redescription. Male. Length 4.4–5.6 mm. General body color black. Antenna, legs and basal part of mandible
dark brown, rest of mandible lighter than basal part (Figs 1, 5, 17).
Head oval, 1.1–1.2 × as broad as high and 1.3–1.5 × as broad as mesosoma, with dense scale-like setae
covering frons from base of mandible to slightly below eye height, scattered scale-like setae reaching top of eye
and behind head; vertex with a few scattered scale-like setae; gena covered or with scattered scale-like setae;
plumose setae long, dark and scattered over vertex and frons; vertex with spaced punctation (Fig. 17). POL 1.0–1.5
× as long as OOL, POL 1.7–2.3 × as long as LOL; eyes ovoid, separated by 0.9–1.0 × their height, malar space
0.03–0.06 × eye height. Scape with scale-like and scattered dark plumose setae, except ventrally glabrous; scape
1.8–2.1 × as long as broad and 2.0–2.6 × as long as pedicel; flagellomeres cylindrical, F1 0.7–0.9 × length of scape
and 0.7–0.8 × length of F2, F2–F6 approximately same length, F7–F10 gradually decreasing in length, F11
elongated with rounded apex.
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FIGURES 1–8. Eotilla spp. 1, E. mickeli Schuster, ♂ (El Canelo, Santiago, Chile), habitus (lateral). 2, E. superba Brothers, ♂
(Posta de Hornillos, Jujuy, Argentina), habitus (lateral). 3, E. medanito sp. nov., ♂ (Saujil, Catamarca, Argentina), habitus
(lateral). 4, E. schusteri sp. nov., ♂ (Udpinango, La Rioja, Argentina), habitus (lateral). 5, E. mickeli Schuster, ♂ (El Canelo,
Santiago, Chile), habitus (dorsal). 6, E. superba Brothers, ♂ (Posta de Hornillos, Jujuy, Argentina), habitus (dorsal). 7, E.
medanito sp. nov., ♂ (Saujil, Catamarca, Argentina), habitus (dorsal). 8, E. schusteri sp. nov., ♂ (Udpinango, La Rioja,
Argentina), habitus (dorsal).
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FIGURES 9–16. Eotilla spp. 9–10, E. mickeli Schuster, ♂ (Perales, Chile): 9, metasoma (dorsal); 10, metasoma (ventral).
11–12, E. superba Brothers, ♂ (Posta de Hornillos, Jujuy, Argentina): 11, metasoma (dorsal); 12, metasoma (ventral). 13–14, E.
medanito sp. nov., ♂ (Saujil, Catamarca, Argentina): 13, metasoma (dorsal); 14, metasoma (ventral). 15–16, E. schusteri sp.
nov., ♂ (Udpinango, La Rioja, Argentina): 15, metasoma (dorsal); 16, metasoma (ventral).
Mesosoma elongate, 1.6–2.1 × as long as broad; punctation well marked, slightly contiguous on dorsum and
more spaced on pleura and vertical surface of propodeum. Pronotum transverse, anteriorly forming a collar with
white and long plumose setae, rest covered almost completely with scale-like and white plumose setae, plumose
setae dark on dorsum. Mesonotum moderately punctate; with scattered scale-like and dark plumose setae, scalelike setae denser on posterior margin; tegula convex, smooth and basal half covered with scale-like setae; scutellum
with scattered scale-like and dark plumose setae. Metanotum transverse, narrow, with scattered scale-like and a few
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dark plumose setae. Propodeum with contiguous punctation; with small dark plumose setae dorsally; vertical
surface with scattered long white plumose setae, except central part smooth and without setae; rest of propodeum
covered with scale-like and white plumose setae; a group of scale-like setae at insertion of hind coxa. Propleuron
covered with plumose setae; mesopleuron convex, punctate except smooth anterodorsal to mid coxa, almost
covered by scale-like and white plumose setae, but less dense medially and above mid coxa exposing the cuticle;
metapleuron with spaced punctation, metepisternum with scattered long plumose setae and with scale-like setae
above insertion of metacoxa, metepimeron covered with scale-like setae on upper half.
Fore wing 2.8–3.1 × as long as broad, densely setose, hyaline, except a narrow small spot posterior to marginal
cell; venation light brown, with coarse dark plumose setae.
Legs slender; anterior surface of coxae, trochanters and femora almost covered by scale-like setae; plumose
setae scattered across surfaces of legs, becoming denser where no scale-like setae; mid and hind tibial spurs white.
Metasoma 2.4–2.6 × as long as broad, T1 1.2–1.7 × as long as wide and 0.6 × as wide as T2 (Fig. 9); terga with
well-marked and contiguous punctation. T1 covered with erect white plumose setae and scattered dark plumose
setae on dorsum, without or with a few scattered scale-like setae on central area, and dense scale-like setae along
posterior margin; T2 entirely covered with erect dark plumose setae and decumbent dark simple setae and scattered
white plumose setae on sides, dense scale-like setae along posterior margin; dark felt line 0.28–0.52 × lateral length
of T2, its anterior extremity at 0.21–0.31 × lateral length of T2; T3–T5 with erect dark plumose setae and
decumbent dark simple setae, scale-like setae on posterior margins; T6 with oblique dark brown coarse plumose
setae without scale-like setae; T7 without a depression, and covered with dark brown coarse plumose setae, shorter
than T6; sterna with sculpture and distribution of setae similar to corresponding terga, except erect plumose setae
white (but becoming dark on posteriormost sterna), S1 without scale-like setae, S6 with scale-like setae on
posterior margin (Fig. 10).
Variation. The number of scale-like setae is variable, but the setation pattern is consistent.
Discussion. Schuster (1949) based his description on seven males from Chile, and Mickel (1968) described the
female based in a (misidentified) specimen from Argentina. There are many differences between species from
Argentina and Chile, and it would be wrong to think that those are conspecific. We believe that Mickel’s female
corresponds to a new species (E. schusteri) described below.
Distribution. Chile: Atacama, Biobío, Copiapó, Coquimbo, Quillota, Maule, Santiago, Valparaíso (Fig. 38).
Phenology. September–February.
Material examined. Paratype of Eotilla mickeli, 1♂: CHILE: Concepción [36°48'47"S 73°02'54"W], 6/XII/
1908, Herbst (1♂, ZMHB). Other material, 9♂: CHILE: Perales, II/1925, J.C. Brethes (no further data) (1♂,
MACN); Coquimbo, Trapiche [30°38'1"S 71°19'50"W], 2/II/1972, Sielfeld (1♂, MNHNS); Valparaíso [33°4'47"S
71°34'49"W], 30/X/1972, Sielfeld (1♂, MNHNS); Olmué [32°59'45"S 71°11'29"W], 21/X/1917, Herbst (1♂,
CAS); Santiago, Caleu [32°59'54"S 70°58'36"W], 6/XI/1988, M. Cerda (1♂, MNHNS); Curacaví [33°24'16"S 71°
7'28"W], 27/IX/1971, H. Sielfeld (1♂, MNHNS); El Alfalfal [33°25'25"S 70°45'11"W], 25/I/1968, J. Moroni (1♂,
MNHNS); El Canelo [33º37'35''S 70º46'35''W], 26/XI/1951, L.E. Peña (1♂, CNC); Maule, Curicó, Huecahuecan
[35°04'15"S 71°09'15"W], 22–28/I/1998, Ugarte (1♂, DJBC).
Eotilla superba Brothers, 1974
(Figs 2, 6, 11, 12, 18, 22, 28, 32, 33)
Eotilla superba Brothers, 1974: 361–363; holotype ♂, Argentina: Catamarca, Nacimientos de Abajo [27°09'54"S 66°43'56"W],
examined.
Diagnosis. Male. Head, mesosoma and metasoma black (Figs 2, 6). T2 with a group of white scale-like setae on
central area (Fig. 11). T7 with an inverted-V-shaped depression medially. Posterior terga and sterna without dark,
short and coarse setae. Female. General body color light brown to yellowish (Figs 22, 28). Posterior surface of
propodeum with transverse striation. Head with dense scale-like setae covering frons from base of mandible to top
of eye; gena without scale-like setae (Fig. 32). T2 and S2 with a group of scale-like setae on each side of posterior
margin (Figs 22, 28). Metanotum with a few scattered scale-like setae, specially on sides dorsally (Fig. 28).
Description. Male. Length 3.7–6.8 mm. General body color black; flagellomeres, tibiae and tarsomeres, and
basal part of mandible dark brown, rest of mandible light brown; rest of antennae and legs black (Figs 2, 6).
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FIGURES 17–21. Eotilla spp. 17, E. mickeli Schuster, ♂ (Perales, Chile), head (anterior). 18, E. superba Brothers, ♂ (Posta de
Hornillos, Jujuy, Argentina), head (anterior). 19, E. schusteri sp. nov., ♂ (Udpinango, La Rioja, Argentina), head (anterior). 20,
E. medanito sp. nov., ♂ (Saujil, Catamarca, Argentina), head (anterior). 21, E. schusteri sp. nov., ♂ (Udpinango, La Rioja,
Argentina), genitalia (ventral; genital foramen projection indicated).
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FIGURES 22–31. Eotilla spp. 22, E. superba Brothers, ♀ (Posta de Hornillos, Jujuy, Argentina), habitus (lateral). 23, E.
medanito sp. nov., ♀ (Udpinango, La Rioja, Argentina), habitus (lateral). 24, E. medanito sp. nov., ♀ (Saujil, Catamarca,
Argentina), habitus (lateral). 25, E. medanito sp. nov., ♀ (Saujil, Catamarca, Argentina), habitus (dorsal). 26, E. schusteri sp.
nov., ♀ (Udpinango, La Rioja, Argentina), habitus (lateral). 27, E. schusteri sp. nov., ♀ (Andalgalá, Catamarca, Argentina),
habitus (lateral). 28, E. superba Brothers, ♀ (Posta de Hornillos, Jujuy, Argentina), habitus (dorsal). 29, E. medanito sp. nov., ♀
(Udpinango, La Rioja, Argentina), habitus (dorsal). 30, E. schusteri sp. nov., ♀ (Udpinango, La Rioja, Argentina), habitus
(dorsal). 31, E. schusteri sp. nov., ♀ (Andalgalá, Catamarca, Argentina), habitus (dorsal).
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FIGURES 32–37. Eotilla spp. 32–33, E. superba Brothers, ♀ (Posta de Hornillos, Jujuy, Argentina): 32, head (anterior); 33,
mesosoma (lateral). 34–35, E. medanito sp. nov., ♀ (Saujil, Catamarca, Argentina): 34, head (anterior); 35, mesosoma (lateral).
36–37, E. schusteri sp. nov., ♀: 36, head (anterior) (Udpinango, La Rioja, Argentina); 37, mesosoma (lateral) (Andalgalá,
Catamarca, Argentina).
Head oval, 1.1–1.4 × as broad as high and 1.2–1.4 × as broad as mesosoma, with dense scale-like setae
covering frons from base of mandible to slightly below eye height, a few scattered scale-like setae reaching top of
eye and behind head; vertex without or with a few scattered scale-like setae; gena covered with scale-like setae;
plumose setae long, dark and scattered over vertex and frons, almost reaching toruli (Fig. 18); vertex with spaced
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punctation. POL 1.5–2.3 × as long as OOL, POL 1.7–2.0 × as long as LOL; eyes ovoid, separated by 0.9–1.1 ×
their height, malar space 0.04–0.09 × eye height. Scape with scale-like and scattered dark plumose setae, except
ventrally glabrous; scape 1.6–2.1 × as long as broad and 1.9–3.3 × as long as pedicel; flagellomeres cylindrical, F1
0.7–0.9 × length of scape and 0.7–0.8 × length of F2, F2–F6 approximately same length, F7–F10 gradually
decreasing in length, F11 elongated with rounded apex.
Mesosoma elongate, 1.5–2.0 × as long as broad; punctation well marked, spaced on dorsum and more
contiguous on pleura and propodeum. Pronotum transverse, anteriorly forming a collar with small plumose setae,
rest covered almost completely with scale-like and white plumose setae, plumose setae dark on dorsum.
Mesonotum moderately punctate; with scattered scale-like and dark plumose setae, scale-like setae denser on
posterior margin; tegula convex, smooth and basal half covered with scale-like setae; scutellum with well-scattered
punctation except medially smooth and with scattered scale-like and dark plumose setae. Metanotum transverse,
narrow, with scattered scale-like and white plumose setae; punctate except medially smooth. Propodeum with
contiguous punctation; with small dark plumose setae dorsally; vertical surface with scattered long white plumose
setae; rest of propodeum covered with scale-like and white plumose setae; a group of scale-like setae at insertion of
hind coxa. Propleuron covered with plumose setae; mesopleuron convex, punctate except smooth anterodorsal to
mid coxa, fairly densely covered by scale-like and white plumose setae, but less dense medially and above mid
coxa exposing the cuticle; metapleuron with moderately spaced punctation but sparser anteroventrally,
metepisternum covered by long plumose setae on central area and with scale-like setae above insertion of
metacoxa, metepimeron covered with scale-like setae.
Fore wing 2.8–3.4 × as long as broad, densely setose (Fig. 2), hyaline, except a narrow small darker spot apical
to marginal cell; venation light brown, with coarse dark plumose setae.
Legs slender; anterior surfaces of coxae, trochanters, femora and tibiae almost covered by scale-like setae;
plumose setae scattered across surfaces of legs, becoming denser where no scale-like setae; mid and hind tibial
spurs white.
Metasoma 2.3–2.8 × as long as broad, T1 1.2–1.5 × as long as wide and 0.6–0.7 × as wide as T2 (Fig. 11); terga
with well-marked punctation, slightly spaced on T1, remaining terga with punctation contiguous. T1 covered with
erect white plumose setae, a few scattered scale-like setae on central area and dense scale-like setae along posterior
margin; T2 with an oval group of few to many scale-like setae on central area, entirely covered with erect white to
dark plumose setae and decumbent dark simple setae, dense scale-like setae along posterior margin; dark felt line
0.37–0.54 × lateral length of T2, its anterior extremity at 0.22–0.30 × lateral length of T2; T3–T5 with scattered
erect white to dark plumose setae and decumbent dark simple setae, scale-like setae on posterior margins; T6 with
scattered erect white to dark plumose setae, apical margin with oblique dark brown coarse plumose setae (different
from those observed in E. mickeli) and a few scale-like setae laterally; T7 with a smooth inverted-V-shaped
depression medially, with scattered small dark plumose to strong simple setae elsewhere; sterna with sculpture and
distribution of setae similar to corresponding terga, except erect plumose setae white (but becoming dark on
posteriormost sterna), S1 without or with only a few scale-like setae posterolaterally, S2 with a few to many
scattered scale-like setae on sides and central area, S6 with only slender scale-like setae on posterior margin, S7
asetose and S8 with dark semi-erect plumose and decumbent strong simple setae (Fig. 12).
Female (hitherto unknown). Length 3.2–4.0 mm. General body color light brown to yellowish except last
segment of metasoma which is dark brown; scape, pedicel, inner surface of flagellomeres and legs light brown;
basal ¾ of F1 black; rest of antenna, apical half of mandible, T6 and S6 dark brown (Figs 22, 28, 32, 33).
Head 1.1–1.3 × as broad as high and 1.3–1.5 × as broad as mesosoma; with dense scale-like setae covering
frons from base of mandible to top of eye; scattered dark plumose setae distributed throughout head, longer on
clypeus; vertex with spaced punctation and coarse dark plumose setae; gena with scattered small dark plumose
setae, without scale-like setae (Figs 32, 33). Eyes separated by 1.1–1.2 × their height, malar space 0.20–0.25 × eye
height. Clypeus short and broad, slightly convex; mandible arched, simple, with small preapical denticle on inner
edge, distance between inner denticle and apical tooth 0.2–0.3 × mandible length. Scape covered with scattered
scale-like and plumose setae on dorsal and inner surfaces, with long plumose setae on ventral surface near insertion
of pedicel, scape 2.0–2.6 × as long as broad and 1.3–1.6 × as long as pedicel; pedicel 2.0–2.7 × as long as broad and
as long as F1; flagellomeres cylindrical, F1 0.6–0.8 × length of scape and 0.8–1.0 × as long as F2; remaining
flagellomeres approximately same length except F6–F9 gradually decreasing in length, F10 elongated with
rounded apex.
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TORRENS ET AL.
Mesosoma elongate, 1.6–2.0 × as long as broad (Fig. 28). Pronotum 1.3–1.6 × as broad as long (width taken
across humeral tubercles), forming a collar anteriorly; humeral tubercle poorly developed; dorsum with wellmarked and contiguous punctation, with a dense transverse band of scale-like setae anteriorly and a few scattered
scale-like setae on posterior margin, dark plumose setae throughout, setae longer laterad; lateral face smooth and
without setae, except a few scale-like setae near insertion of fore coxa. Fused metanotum and propodeum dorsally
with scattered and well-marked punctation, with a few scattered scale-like setae, especially on sides, and scattered
dark plumose setae which longer laterad; posterior declivity of propodeum without setae and transversely striate,
striations extending beyond limit of metapleura, sometimes reaching to level of spiracle. Propleuron smooth, with
scattered dark plumose setae and a group of dark long plumose setae near insertion of fore coxa; mesopleuron
smooth with a longitudinal carina reaching slightly beyond half of sclerite from ventral limit of pronotum, with
scattered scale-like and small plumose setae, scale-like setae denser above mid coxa; metapleural-propodeal suture
indistinguishable, without setae except a few scale-like setae near insertion of mid coxa and above hind coxa (Fig.
33).
Legs slender; fore coxa with a group of dark long plumose setae on upper outer surface and a few scale-like
and long plumose setae elsewhere; mid and hind coxae, trochanters, femora and tibiae covered with scale-like and
plumose setae, except no scale-like setae on ventral surfaces.
Metasoma 1.9–2.2 × as long as broad (Fig. 28), T1 0.93–0.96 × as long as wide and 0.52–0.60 × as wide as T2;
with well-marked contiguous punctation on T1 and T2, except central area of vertical surface of T1 smooth; T3–T5
with punctation less marked and dispersed; T6 completely alutaceous. T1 with dense group of scale-like setae
laterally on dorsal surface, scattered dark long plumose setae except medially on vertical surface; T2 with dense
group of scale-like setae posterolaterally, with dark plumose setae across surface, dark felt line 0.14–0.17× lateral
length of T2, its anterior extremity 0.24–0.26 × lateral length of T2; T3 with dark plumose setae; T4–T5 with
scattered dark plumose setae on posterior margin; T6 without setae. Sterna with sculpture and distribution of setae
similar to that of corresponding terga, except S2 with a few scale-like setae on each side of posterior margin, and
S6 with latero-posterior margin with distinct denticles.
Variation. Males. In some specimens the coloration of the metasoma and/or legs, particularly the tibiae and
tarsomeres, is lighter. The number of scale-like setae is variable, specially on T1, T2 and S2; but the setation
pattern is rather consistent.
Females. In some specimens the coloration of the legs is darker, particularly the fore and mid coxae, femora
and tibiae, and the metasoma a little lighter than the rest of the body. The number of scale-like setae is variable, but
the setation pattern is consistent. In some specimens the striation of the propodeum is weak, appearing smooth.
Discussion. The association between males and females is very difficult because of extreme sexual
dimorphism. However, several specimens of both sexes were collected in the same place and on the same day, in
addition to being the only species found at that locality, so that they are considered conspecific.
Distribution. Argentina: Catamarca, Jujuy, La Rioja, Salta (Fig. 38).
Phenology. September–March.
Material examined. Holotype of Eotilla superba, ♂ : ARGENTINA: Catamarca, Los Nacimientos de Abajo
[27°10'42"S 66°43'34"W], 2060 m, 16–31/XII/1968, Willink, Terán and Stange, MT (IFML). Paratypes, 4♂:
ARGENTINA: Salta, Yacochuya (Cafayate) [26°2'16"S 66°1'29"W], 1950 m, 1–15/IX/1968, Willink, Terán and
Stange, MT (2♂, IFML); same location, 16–30/IX/1968, Willink, Terán and Stange, MT (1♂, DJBC); Salta, Tacuil
[25°35'15"S 66°26'23"W], 7/XII/1968, Willink, Stange (1♂, DJBC). Other material, 66♂ and 9♀: Catamarca, 10
km S of Cortaderas 27º37'57''S 68º14'52''W, 3923 m, 18/XII/2012, J. Torréns and P. Fidalgo, YPT (1♂, IFML);
Jujuy, Ruta 9, Km 1762, ca Posta de Hornillos 23°39'47"S 65°26'13"W, 2390 m, 2–4/IX/2009, P. Fidalgo, YPT (12
♂, 1♀, MACN); same location, 18–20/I/2012, J. Torréns and P. Fidalgo, YPT (16♂, 6♀, IFML); Ruta 52, Km 17,
Puerta de Lipán 23°40'01"S 65°34'46"W, 2954 m, 10–11/X/09, P. Fidalgo, YPT (9♂, IADIZA); same location,
18–20/I/2012, J. Torréns and P. Fidalgo, YPT (1♀, DJBC); Ruta 52, Km 18, Puerta de Lipán 23°40'1"S
65°35'11"W, 3008 m, 10–11/X/09, P. Fidalgo, YPT (10♂, DJBC; 10♂, MNHNS); same location, 18–20/I/2012, J.
Torréns and P. Fidalgo, YPT (2♂, 1♀, MNHNS); Quera 23°5'58"S 65°46'11"W, 5–6/III/2013, 3500 m, J. Torréns
and P. Fidalgo, YPT (1♂, IFML); La Rioja, El Carrizal, 28°54'35"S 66°39'26"W, 16–18/XI/09, A. Aranda, pit-fall
(1♂, MACN); La Puerta, 28°49'39"S 66°39'20"W, 29/XI/09, Torréns and Fidalgo, YPT (2♂, MACN); Qda. Santo
Domingo 28°33'18"S 68°44'46"W, 2900 m, 02/XI/2009, J. Torréns, YPT (2♂, IFML).
REVIEW OF THE GENUS EOTILLA SCHUSTER, 1949
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FIGURE 38. Distribution of species of Eotilla in Argentina and Chile.
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TORRENS ET AL.
Eotilla medanito Torréns, Fidalgo, Roig-Alsina & Brothers, sp. nov.
(Figs 3, 7, 13, 14, 20, 23–25, 29, 34, 35)
Diagnosis. Male. Head and mesosoma black (Figs 3, 7), metasoma entirely orange but apical terga darkened (Figs
13, 14). T2 with a group of scale-like setae on central area. Posterior terga and sterna without dark, short and coarse
setae. T7 with an inverted-V-shaped depression medially. Female. General body color light brown to reddish,
sometimes pleura, propodeum, legs and last two or three segments of metasoma with dark brown area (Figs 23–25,
29). All terga and sterna, except T1, without scale-like setae (Figs 23, 25, 29). Posterior surface of propodeum with
striation, especially well marked on sides.
Description. Male. Length 4.3–4.7 mm. Head and mesosoma black, metasoma orange. Scape, pedicel,
flagellomeres except outer surface, coxae, fore and mid femora and tibia dark brown; rest of legs and antennae light
brown. (Figs 3, 7).
Head oval, 1.1–1.3 × as broad as high and 1.2–1.4 × as broad as mesosoma, with dense scale-like setae
covering entire head, except around ocelli; gena covered with scale-like setae; plumose setae long, white and
scattered over entire head, including vertex and occiput; vertex smooth (Fig. 20). POL 1.7–1.9 × as long as OOL,
POL 1.6–2.1 × as long as LOL; eyes ovoid, separated by 1.0–1.1 × their height, malar space 0.05–0.13 × eye
height. Scape with scale-like and white plumose setae covering almost entire surface except longitudinal carina
ventrally; scape 1.8–1.9 × as long as broad and 1.9–2.6 × as long as pedicel; flagellomeres cylindrical, F1 0.8–0.9 ×
length of scape and 0.7–0.8 × length of F2, flagellomeres F2–F4 approximately same length, rest of flagellomeres
gradually decreasing in length, F11 elongated with rounded apex.
Mesosoma elongate, 1.6–1.9 × as long as broad (Fig. 7); punctation well marked, spaced on dorsum and more
contiguous on pleura and propodeum. Pronotum transverse, anteriorly forming a collar with small plumose setae,
rest covered completely with scale-like setae and scattered white plumose setae. Mesonotum moderately punctuate;
with scattered scale-like and white plumose setae, scale-like setae denser on posterior margin; tegula convex,
smooth and covered with scale-like setae; scutellum with scale-like and white plumose setae covering the entire
surface. Metanotum transverse, narrow, with scale-like and white plumose setae. Propodeum with contiguous
punctation becoming spaced on vertical surface; with small dark plumose setae dorsally; vertical surface with a few
scale-like setae and with scattered long white plumose setae; rest of propodeum covered with scale-like and white
plumose setae. Propleuron covered almost completely with scale-like setae and long white plumose setae;
mesopleuron convex, densely covered by scale-like and plumose setae; metapleuron with spaced punctation,
metepisternum covered by long plumose setae on central area and with scale-like setae at insertion of metacoxa,
metepimeron covered with scale-like setae.
Fore wing 3.2–3.4 × as long as broad, moderately setose (Fig. 3), hyaline; venation light brown, with coarse
dark plumose setae on costal and subcostal veins.
Legs slender; anterior surface of coxae, trochanters, femora and tibiae covered by scale-like setae; plumose
setae scattered across surfaces of legs, becoming denser where no scale-like setae; mid and hind tibial spurs white.
Metasoma 2.4–2.6 × as long as broad, T1 1.2–1.5 × as long as wide and 0.6 × as wide as T2 (Fig. 13); terga
with well-marked contiguous punctation. T1 covered with scale-like setae, except central area dorsally with
plumose setae; T2 with a group of scale-like setae on antero-central area and posterior margin, elsewhere with
scattered white plumose setae and decumbent dark simple setae; white felt line 0.38–0.50 × lateral length of T2, its
anterior extremity at 0.17–0.28 × lateral length of T2; T3 with white plumose setae and decumbent dark simple
setae, scale-like setae on posterior margin; T4–T6 covered by scale-like setae and white plumose setae; T6 with
scattered dark plumose setae and oblique dark brown coarse plumose setae on central area of apical margin; T7
with a smooth inverted-V-shaped depression medially, scale-like setae on both sides of depression anteriorly, and
scattered dark brown plumose setae; sterna with sculpture and distribution of setae similar to corresponding terga,
except S1 without or with only a few scale-like setae posterolaterally, S2 with several scattered scale-like setae
covering whole surface, S7 asetose and S8 with dark semi-erect plumose and decumbent strong simple setae and a
few scale-like setae (Fig. 14).
Female. Length 2.7–3.7 mm. General body color light brown to reddish (Figs 23, 29), sometimes pleura,
propodeum and last two or three segments of metasoma with dark brown patch (Figs 24, 25); scape, pedicel and
inner surface of flagellomeres light brown, elsewhere flagellomeres dark brown, basal ¾ of F1 black; apical half of
mandible, coxae and femora darker than rest of body (Figs 23–25, 29, 34, 35).
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Head 1.1–1.3 × as broad as high and 1.3–1.4 × as broad as mesosoma; with dense scale-like setae covering
entire head; scattered dark plumose setae distributed throughout head, longer on clypeus; vertex with spaced
punctation and white plumose setae near occiput; gena covered with scale-like setae and scattered dark plumose
setae (Figs 34, 35). Eyes separated by 1.1–1.3 × their height, malar space 0.24–0.33 × eye height. Clypeus short
and broad, slightly convex; mandible arched, simple, with small preapical denticle on inner edge, distance from
inner denticle to apical denticle 0.22 × mandible length. Scape covered with scale-like and plumose setae on dorsal
and inner surfaces, with long plumose setae on ventral surface near insertion of pedicel, scape 2.0–2.5 × as long as
broad and 1.4–1.7 × as long as pedicel; pedicel 1.8–2.3 × as long as broad; flagellomeres cylindrical, F1 0.5–0.7 ×
length of scape and 1.0 × as long as F2; remaining flagellomeres approximately same length except F6–F9
gradually decreasing in length, F10 elongated with rounded apex.
Mesosoma elongate, 1.5–1.9 × as long as broad (Figs 25, 29). Pronotum 1.4–1.7 × as broad (across humeral
tubercles) as long (excluding anterior collar); humeral tubercle poorly developed; dorsum with well-marked and
contiguous punctation anteriorly and spaced punctation on central area, covered with scale-like setae and scattered
dark plumose setae throughout, setae longer laterad; lateral face covered with scale-like setae. Fused metanotum
and propodeum dorsally covered with scale-like setae and scattered dark plumose setae throughout, plumose setae
longer laterad; posterior declivity of propodeum smooth on upper half and striate on lower half especially well
marked on sides, with scattered small plumose setae and a few scale-like setae laterad, reaching to insertion of hind
coxa. Propleuron smooth and without setae, except a group of dark long plumose setae near insertion of fore coxa;
mesopleuron weakly rugose to smooth with a longitudinal carina reaching to half of sclerite from ventral limit of
pronotum, covered with scale-like setae, denser above mid coxa and long plumose setae at insertion of mid coxa;
metapleural-propodeal suture indistinguishable, with scattered plumose setae and a longitudinal band of scale-like
setae above hind coxa continuous with that on mesopleuron, reaching propodeal foramen and connected with scalelike setae of propodeum (Fig. 35).
Legs slender; fore coxa transversely striate, with a group of dark long plumose setae on upper outer surface and
distributed at random near insertion of trochanter and on inner surface and few small scale-like setae on outer and
posterior surfaces and near insertion of trochanter; mid and hind coxae, trochanters, femora and tibiae covered with
scale-like and plumose setae, except no scale-like setae on ventral surfaces.
Metasoma 2.0–2.4 × as long as broad (Figs 25, 29); T1 0.84–0.93 × as long as wide and 0.60–0.70 × as wide as
T2; with well-marked contiguous punctation on T1, except central area of vertical surface of T1 smooth; T2–T5
with punctation less marked and dispersed; T6 completely alutaceous. T1 covered with scale-like and scattered
dark long plumose setae, except vertical surface without scale-like setae and with a few white long plumose setae;
T2 with dark plumose setae across surface, dark felt line 0.17–0.20× lateral length of T2, its anterior extremity
0.20–0.26 × lateral length of T2; T3–T5 with scattered dark plumose setae; T6 without setae. Sterna with sculpture
and distribution of setae similar to that of corresponding terga, except S1 with a few scattered plumose setae and
without scale-like setae, and S6 with latero-posterior margin with distinct denticles.
Variation. Males. The antennae and legs are sometimes darker, and the femora may be almost completely dark
brown; the predominance of scale-like setae on the mesosoma is higher in some specimens; on T2 the scale-like
setae may cover almost the entire dorsal region.
Females. The general body color varies among individuals, being slightly darker in some specimens,
especially from Saujil (Figs 24, 25), however the general color pattern is retained. Some specimens have fewer
scale-like setae on the dorsum of the pronotum. In some specimens the upper half of the propodeum is rugose, and
the striation laterally reaching to the dorsal surface.
Discussion. As for E. superba, several specimens of both sexes were collected at the same place and on the
same day, in addition to being the only species found there, so that they are considered conspecific.
Etymology. The name “medanito” comes from the Spanish diminutive of “médano” and refers to the dune
environment where this species lives. It is a noun in apposition.
Distribution. Argentina: Catamarca, La Rioja, Santa Cruz (Fig. 38).
Phenology. November–May.
Material examined. Holotype ♂ : ARGENTINA: Catamarca, 3.5 km N de Saujil, Ruta 34 27°32'06"S
67°33'23"W, 1650 m, 4–6/XII/2011, P. Fidalgo, YPT (MACN). Paratypes, 5♂ and 23♀: Catamarca, 3.5 km N de
Saujil, Ruta 34 27°32'06"S 67°33'23"W, 1650 m, 4–6/XII/2011, P. Fidalgo, YPT (1♂, IFML); same location, 8/II/
2006, P. Fidalgo and P. Diez, YPT (3♀, IFML); same location, 13–14/xi/2013, Fidalgo and Torréns (11♀, MACN);
14 · Zootaxa 3878 (1) © 2014 Magnolia Press
TORRENS ET AL.
same location, 1/v/2014, P. Fidalgo (1♂, 2♀, DJBC); La Rioja, sur de Udpinango 28°42'7"S 66°47'51"W, 991 m,
II/2006, P. Fidalgo, G. Fidalgo, P. Diez and P. Torréns, YPT (4♀, IFML); same location, 25/II–4/III/2006, Fidalgo,
Diez and Torréns, pit-fall (1♀, DJBC); same location, 20/III/2006, Fidalgo, Diez and Torréns, pit-fall (1♀, DJBC);
same location, 8–28/IV/2006, pit-fall (1♀, DJBC); same location, I/2007, J. Torréns and C. Nieto, YPT (2♂,
MACN); same location, 2–9/XI/2011, J. Torréns and P. Fidalgo, YPT (1♂, 1♀, IFML); Santa Cruz, 2 km S of
Caleta Olivia [46°29'36"S 67°32'55"W], 12/XII/1966, 10 m, E.I. Schlinger and M. Irwin (1♂, CAS).
Eotilla schusteri Torréns, Fidalgo, Roig-Alsina & Brothers, sp. nov.
(Figs 4, 8, 15, 16, 19, 21, 26, 27, 30, 31, 36, 37)
Eotilla mickeli female, Mickel 1968: 234–236 (misidentification).
Diagnosis. Male. Head, mesosoma and metasoma dark brown to black (Figs 4, 8). T1, T2 and S2 with abundant
scale-like setae on all surfaces (Figs 15, 16). Female. Head and pronotum brownish to reddish; metasoma entirely
dark brown to black or T1 and S1 brownish to reddish and remainder darker (Figs 26, 27, 30, 31). T2 with abudant
scale-like setae, except on central area (Figs 30, 31). Posterior surface of propodeum with striation, especially well
marked on sides.
Description. Male. Length 4.0–5.7 mm. General body color black. Flagellomeres, femora, tibiae and
tarsomeres, and apical part of mandible dark brown; scape and pedicel dark brown to black; rest of mandible light
brown (Figs 4, 8).
Head oval, 1.0–1.2 × as broad as high and 1.2–1.4 × as broad as mesosoma, with dense scale-like setae
covering frons from base of mandible to top of eye, a few scattered scale-like setae on vertex and behind head;
gena covered with scale-like setae; plumose setae long, white and scattered over vertex and frons and with a few
dark plumose setae on vertex (Fig. 19); vertex with spaced punctation. POL 1.8–2.1 × as long as OOL, POL
1.7–2.2 × as long as LOL; eyes ovoid, separated by 1.0–1.1 × their height, malar space 0.04–0.09 × eye height.
Scape with scale-like and scattered white plumose setae, except ventrally glabrous; scape 1.6–2.1 × as long as
broad and 2.0–2.8 × as long as pedicel; flagellomeres cylindrical, F1 0.8 × length of scape and 0.6–0.8 × length of
F2, F2–F6 approximately same length, F7–F10 gradually decreasing in length, F11 elongated with rounded apex.
Mesosoma elongate, 1.6–1.9 × as long as broad; punctation well marked and contiguous, spaced on dorsum of
mesonotum and scutellum. Pronotum transverse, anteriorly forming a collar with white scattered plumose setae,
rest covered completely with scale-like and white plumose setae and a few dark plumose setae. Mesonotum
moderately punctate; with scattered scale-like, white plumose setae and a few dark plumose setae, scale-like setae
denser on posterior half; tegula convex, smooth and covered with scale-like setae; scutellum with scattered
punctation and with scattered scale-like and white plumose setae and a few dark plumose setae. Metanotum
transverse, narrow, almost covered with scale-like and white plumose setae; punctate. Propodeum with contiguous
punctation; with small dark plumose setae dorsally; vertical surface with scattered long white plumose setae; rest of
propodeum covered with scale-like and white plumose setae. Propleuron covered with long scale-like and plumose
setae; mesopleuron convex, punctate, fairly densely covered by scale-like and white plumose setae; metapleuron
with moderately spaced punctation but sparser anteroventrally, metepisternum covered by long plumose setae on
central area and with scale-like setae around central area and above insertion of metacoxa, metepimeron covered
with scale-like setae.
Fore wing 2.9–3.4 × as long as broad, densely setose (Fig. 4), hyaline; venation light brown, with coarse dark
plumose setae.
Legs slender; anterior surfaces of coxae, trochanters, femora and tibiae covered by scale-like setae; plumose
setae scattered across surfaces of legs, becoming denser where no scale-like setae; mid and hind tibial spurs white.
Metasoma 2.3–2.8 × as long as broad, T1 1.1–1.4 × as long as wide and 0.6–0.7 × as wide as T2 (Fig. 15); terga
with well-marked punctation, slightly spaced on T1, remaining terga with punctation contiguous. T1 covered with
erect white plumose setae, with abundant scale-like setae, less dense on central area on dorsum, without scale-like
setae on vertical surface; T2 with a dense group of scale-like setae on anterior margin and reaching to central area,
entirely covered with erect white to dark plumose setae and decumbent dark simple setae, dense scale-like setae
along posterior margin; dark felt line 0.45–0.57 × lateral length of T2, its anterior extremity at 0.17–0.22 × lateral
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length of T2; T3–T4 with erect white to dark plumose setae and decumbent dark simple setae, scale-like setae on
posterior margins; T5–T6 with scattered scale-like setae and erect white to dark plumose setae, apical margin with
oblique dark brown coarse plumose setae (different from those observed in E. mickeli); T7 with a smooth invertedV-shaped depression medially, with scattered small dark plumose to strong simple setae elsewhere; sterna with
sculpture and distribution of setae similar to corresponding terga, S1 without or with only a few scale-like setae
posterolaterally, S2 almost covered with scattered scale-like setae, S7–S8 with dark semi-erect plumose and
decumbent strong simple setae (Fig. 16).
Female. Length 2.4–4.0 mm. Head, scape, pedicel, apical part of F1 and inner surfaces of flagellomeres,
mesosoma, legs, T1 and S1 brownish to reddish; rest of flagellomeres except basal 3/4 of F1 dark brown; rest of F1
and metasoma brownish to blackish (Figs 26, 27). Some specimens with head and pronotum brownish to reddish
and rest of mesosoma and metasoma dark brown to black (Figs 27, 31).
Head 1.1–1.3 × as broad as high and 1.3–1.4 × as broad as mesosoma; with dense scale-like setae covering
frons from base of mandible to top of eye; scattered dark plumose setae distributed throughout head, longer on
clypeus; vertex with spaced punctation but smooth medially, with a few scale-like setae and coarse dark plumose
setae; gena with scattered scale-like setae covering all surface, without plumose setae (Fig. 36). Eyes separated by
1.0–1.2 × their height; malar space 0.16–0.30 × eye height. Clypeus short and broad, slightly convex; mandible
arched, simple, with small preapical denticle on inner edge, distance from inner denticle to apical denticle
0.14–0.27 × mandible length. Scape covered with scale-like setae on dorsal and inner surfaces, with a few short
plumose setae near insertion of pedicel and long plumose setae on ventral surface, scape 2.4–3.1 × as long as broad
and 1.3–2.2 × as long as pedicel; pedicel 1.8–2.2 × as long as broad; flagellomeres cylindrical, F1 0.5–0.7 × length
of scape and as long as F2; remaining flagellomeres approximately same length, except F5–F9 gradually
decreasing in length, F10 elongated with rounded apex.
Mesosoma elongate, 1.8–2.1 × as long as broad (Figs 30, 31). Pronotum 1.1–1.6 × as broad as long (across
humeral tubercles) as long (excluding anterior collar); humeral tubercle poorly developed; dorsum with wellmarked and very spaced punctation, with scale-like setae covering entire surface or restricted to a transverse band
anteriorly, dark plumose setae throughout, setae longer laterad; lateral surface punctate or smooth, with scale-like
setae covering entire surface or scattered and denser above fore coxa. Fused metanotum and propodeum dorsally
with spaced and well-marked punctation becoming contiguous on sides, with scale-like setae covering dorsal
surface and scattered dark plumose setae throughout, setae longer laterad; posterior declivity of propodeum without
setae and transversely striate; lateral face smooth on upper half and striate on lower half, especially well marked on
sides. Propleuron smooth, with scattered scale-like setae and a group of white or dark long plumose setae near
insertion of fore coxa; mesopleuron smooth with a longitudinal carina restricted to half of sclerite from ventral
limit of pronotum, with dense or scattered scale-like setae denser above mid coxa; metapleural-propodeal suture
indistinguishable, with scattered scale-like setae or with scale-like setae only near insertion of mid coxa and above
hind coxa (Fig. 37).
Legs slender; fore coxa with a group of white or dark long plumose setae on upper outer surface and
distributed at random in all surfaces, with scattered scale-like and dark long plumose setae elsewhere; mid and hind
coxae, trochanters, femora and tibiae covered with scale-like and plumose setae, except no scale-like setae on
ventral surfaces.
Metasoma 2.1–2.5 × as long as broad (Figs 30, 31); T1 0.92–1.08 × as long as wide and 0.65–0.75 × as wide as
T2; with well-marked contiguous punctation on T1 and T2, except vertical surface of T1 smooth; T3–T5 with
punctation less marked and dispersed; T6 completely alutaceous. T1 with scale-like setae except central area on
dorsal surface, scattered dark long plumose setae except with white long plumose setae on vertical surface; T2 with
scale-like setae anteriorly and laterally, denser postero-laterally, with dark plumose setae across surface, dark felt
line 0.11–0.20× lateral length of T2, its anterior extremity 0.24–0.32 × lateral length of T2; T3 with a few scale-like
setae posterolaterally or without such setae; T3–T5 with dark plumose setae; T6 without setae. Sterna with
sculpture and distribution of setae similar to that of corresponding terga, except S1 and S2 with a few scale-like
setae, and S6 with latero-posterior margin with distinct denticles.
Variation. Males. In some specimens the coloration of the metasoma and legs, particularly the tibiae and
tarsomeres, is lighter. The number of scale-like setae is variable, specially on T1 and T2; but the setation pattern is
fairly consistent.
Females. The coloration of the mesosoma and metasoma is highly variable; there are specimens with the
16 · Zootaxa 3878 (1) © 2014 Magnolia Press
TORRENS ET AL.
pronotum lighter than the rest of the mesosoma and metasoma, and others with the mesosoma and T1 lighter than
the rest of the metasoma; also, in some specimens the coloration of the head, femora, tibiae, pleura and propodeum
is darker. One specimen from Neuquén (Las Lajas) has a particular coloration of the metasoma, with the vertical
surface of T1, the anterior half of T2 and S2 and the last three terga of the metasoma dark brown and the rest
lighter, but the morphological characters and disposition of setae correspond to this species. The pronotum has a
few scale-like setae on the central area of the dorsum or none; otherwise the scale-like setae on the mesosoma and
metasoma show variation in density but not in disposition.
Discussion. As discussed above for E. superba, all females were collected at the same locality as males, so
they are considered conspecific.
Mickel (1968) described a female specimen from Argentina and attributed it to E. mickeli Schuster (from
Chile); however, the notable differences between species from Argentina and Chile, and the remarkable similarity
of that specimen with the description above, suggests that it was misidentified and is actually this species.
Etymology. Named in honor of the describer of the genus Eotilla (and Prototilla), Rudolf M. Schuster.
Distribution. Argentina: Catamarca, La Rioja, Neuquén (Fig. 38).
Phenology. September, December–April.
Material examined. Holotype ♂: ARGENTINA: La Rioja, Santa Teresita [28°35'57"S 66°33'32"W], 690 m,
8–28/IV/2006, Porter, Torréns y Fidalgo, MT (1♂, MACN). Paratypes, 9♂ and 8♀: Catamarca, Andalgalá
[27°34'60"S 66°21'2"W], 12/XII/1973, JH Hunt #1495, det. as E. mickeli by Snelling (1♂, LACM); Andalgalá,
Villavil [27°34'36"S 66°15'02"W], 30/XII/1973, FE Enders, JHHunt#1504 (2♀ LACM, 1♀ DJBC); same data but
JHH#1502 (1♀ LACM); El Pucará empalme [27°42'17"S 66°00'36"W], 7/I/1974, FE Enders, #JHH1499 det. as E.
mickeli by Snelling (1♀, LACM); La Rioja, Udpinango, 5 km S de Udpinango 28°43'16"S 66°47'34"W, 1011 m,
16–25/III/2005, P. Fidalgo, G. Fidalgo and J. Torréns, YPT (1♀, DJBC); same location, II/2006, Fidalgo, Diez and
Torréns, YPT (1♀, MACN); same location, 8–28/IV/2006, pit-fall (1♀, IFML); same location, 26/I/2007, Torréns,
Fidalgo and Diez, YPT (5♂, MACN); same location, I/2007, J. Torréns and C. Nieto, YPT (1♂, DJBC); same
location, 26/II/2007, Fidalgo, Diez and Torréns, YPT (1♂, IFML); same location, 2–9/XI/2011, J. Torréns and P.
Fidalgo, YPT (1♀, IFML); La Puerta, 28°50'13"S 66°39'29"W, 16–18/XI/09, A. Aranda, pit-fall (1♀, MACN); 7
km E de Anillaco [28°46'29"S 66°51'53"W], IX–X/2013, A. Aranda, pit-fall (1♂, IFML); Neuquén, Covunco
[38°46'1"S 70°3'18"W], 18/I/1949 (Allotype, 1♀, IFML); 20 km Las Lajas, ANP Cuchillo Curá [38°36'40"S
70°23'54"W], 20/I/1992, J. Sganga (1♀, IFML)
Acknowledgements
This work was made possible by funds provided by Agencia Nacional de Promoción Científica y Tecnológica,
PICT 2007-1238, Préstamo BID. The University of KwaZulu-Natal Research Office provided funding to DJB.
Thanks also go to James Pitts and Emily Sadler for their contributions and suggestions to this manuscript, and to
Mario Elgueta (MNHNS) and to the curators of other collections named above for access to specimens in their
care. We thank two anonymous reviewers for the useful suggestions.
References
Ashmead, W.H. (1899) Superfamilies in the Hymenoptera and generic synopses of the families Thynnidae, Myrmosidae, and
Mutillidae. Journal of the New York Entomological Society, 7, 45–60.
Blake, C.A. (1886) Monograph of the Mutillidae of North America. Transactions of the American Entomological Society, 13,
179–286.
Brothers, D.J. (1970) Discovery of the mutillid subfamily Typhoctinae in South America, with description of a new species
(Hymenoptera). Journal of the Kansas Entomological Society, 43, 302–308.
Brothers, D.J. (1974) The new genus Typhoctoides and a new species of Eotilla (Hymenoptera: Bradynobaenidae;
Typhoctinae). Journal of the Kansas Entomological Society, 47, 359–363.
Brothers, D.J. (1975) Phylogeny and classification of the aculeate Hymenoptera, with special reference to Mutillidae. The
University of Kansas Science Bulletin, 50, 483–648.
Brothers, D.J. (1999) Phylogeny and evolution of wasps , ants and bees (Hymenoptera , Chrysidoidea, Vespoidea and
Apoidea). Zoologica Scripta, 28, 233–249.
REVIEW OF THE GENUS EOTILLA SCHUSTER, 1949
Zootaxa 3878 (1) © 2014 Magnolia Press ·
17
http://dx.doi.org/10.1046/j.1463-6409.1999.00003.x
Brothers, D.J. (2006) Capitulo 52, Familia Bradynobaenidae. In: Fernández, F. & Sharkey, M.J. (Eds.), Introducción a los
Hymenoptera de la Región Neotropical. Bogotá, D.C., pp. 559–562.
Brothers, D.J. & Carpenter, J.M. (1993) Phylogeny of Aculeata: Chrysidoidea and Vespoidea (Hymenoptera). Journal of
Hymenoptera Research, 2, 227–304.
Debevec, A.H., Cardinal, S. & Danforth, B.N. (2012) Identifying the sister group to the bees: a molecular phylogeny of
Aculeata with an emphasis on the superfamily Apoidea. Zoologica Scripta, 41, 527–535.
http://dx.doi.org/10.1111/j.1463-6409.2012.00549.x
Eady, R.D. (1968) Some illustrations of microsculpture in the Hymenoptera. Proceedings of the Royal Entomological Society of
London , 43 (A), 66–72.
http://dx.doi.org/10.1111/j.1365-3032.1968.tb01029.x
Fernández, F. (2001) Checklist of Genera and Subgenera of Aculeate Hymenoptera of the Neotropical Region. Biota
Colombiana, 2, 87–130.
Fernández, F. (2002) Filogenia y Sistemática de los Himenópteros con Aguijón de la Región Neotropical (Hymenoptera:
Vespomorpha). In: Costa, C., Vanin, S.A., Lobo, J.M. & Melic, A. (Eds.), Proyecto de Red Iberoamericana de
Biogeografía y Entomología Sistemática PrIBES 2002, m3m: Vol. 2. Monografías Tercer Milenio, pp. 101–138.
Hadley, A. (2010) CombineZP public domain image processing software. Available from: http://
www.hadleyweb.pwp.blueyonder.co.uk/CZP/News.htm (accessed on 10 October 2014)
Harris, R.A. (1979) A glossary of surface sculpturing. Occasional Papers in Entomology, 28, 1–31.
Heraty, J., Ronquist, F., Carpenter, J.M., Hawks, D., Schulmeister, S., Dowling, A.P., Murray, D., Munro, J., Wheeler, W.C.,
Schiff, N. & Sharkey, M. (2011) Evolution of the hymenopteran megaradiation. Molecular Phylogenetics and Evolution,
60, 73–88.
http://dx.doi.org/10.1016/j.ympev.2011.04.003
Krombein, K.V. (1942) A new genus and species of Brachycistidinae (Hymenoptera: Tiphiidae). Proceedings of the
Entomological Society of Washington, 44, 65–68.
Martinez, A. & Fritz, M.A. (1974) Notas sobre Mutillidae III (Hymenoptera). Revista Chilena de Entomología, 8, 23–26.
Mickel, C.E. (1968) The female sex of the Mutillidae subfamily Eotillinae (Hymenoptera). Psyche, 74, 234–236.
Pilgrim, E.M., von Dohlen, C.D. & Pitts, J.P. (2008) Molecular phylogenetics of Vespoidea indicate paraphyly of the
superfamily and novel relationships of its component families and subfamilies. Zoologica Scripta, 37, 539–560.
http://dx.doi.org/10.1111/j.1463-6409.2008.00340.x
Reid, J.A. (1941) The thorax of the wingless and short-winged Hymenoptera. Transactions of the Royal Entomological Society
of London, 91, 367–446.
http://dx.doi.org/10.1111/j.1365-2311.1941.tb01046.x
Schuster, R.M. (1949) Contributions toward a monograph of the Mutillidae of the Neotropical Region. III. A key to the
subfamilies represented and descriptions of several new genera (Hymenoptera). Entomologica Americana, 29, 59–140.
Shorthouse, D.P. (2010) SimpleMappr, an online tool to produce publication-quality point maps. Available from: http://
www.simplemappr.net/ ( accessed on 10 October 2014)
Torréns, J., Fidalgo, P., Roig-Alsina, A. & Brothers, D.J. (2012) New species of Prototilla Schuster, 1949 from Argentina and
diagnoses of the genus based on male and female (Hymenoptera: Bradynobaenidae: Typhoctinae: Eotillini). Zootaxa, 32,
17–32.
Torréns, J. & Roig-Alsina, A. (2009) Description of a new species of Bradynobaenus (Hymenoptera: Bradynobaenidae) from
Argentina, with a key to the females of the genus. Zootaxa, 2047, 63–68.
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TORRENS ET AL.