FRFossil RecordFRFoss. Rec.2193-0074Copernicus GmbHGöttingen, Germany10.5194/fr-18-119-2015The first Stylogaster Macquart, 1835 (Diptera:
Conopidae) fossil, from Oligo-Miocene Dominican amber, and some phylogenetic
and biogeographic considerationsRochaL. S. G.leonardo.rocha@ifrj.edu.brBurtT. O.de Mello-PatiuC. A.SkevingtonJ. H.Graduate Program in Biological Sciences – Zoology, Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro, RJ, BrazilInstituto Federal de Educação, Ciência e Tecnologia do Rio de Janeiro, Campus Nilópolis, Nilópolis, RJ, BrazilCanadian National Collection of Insects, Arachnids and Nematodes, Agriculture and Agri-Food Canada, Ottawa, ON, CanadaDepartment of Biology, Carleton University, Ottawa, ON, CanadaDepartment of Entomology, Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro, RJ, BrazilResearch fellow of Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), Brazilnow at: Department of Entomology, Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro, RJ, BrazilL. S. G. Rocha (leonardo.rocha@ifrj.edu.br)29June20151821191255April201514May201518May2015This work is licensed under a Creative Commons Attribution 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by/3.0/This article is available from https://fr.copernicus.org/articles/18/119/2015/fr-18-119-2015.htmlThe full text article is available as a PDF file from https://fr.copernicus.org/articles/18/119/2015/fr-18-119-2015.pdf
Stylogaster Macquart, 1835 has been unknown in the fossil record until now, the only
fossil conopid genus being Palaeomyopa Meulnier, 1912. Two Stylogaster specimens in amber from the
American Museum of Natural History collection were studied and are described
here. Both specimens, male and female, belong to a new species, S. grimaldii sp. nov.,
that is probably basal to at least the New World species with a short
ocellar triangle. Photos and drawings of the new species are provided. The
relationship with other Stylogaster species and ancient distributional patterns are
briefly discussed.
Introduction
Conopidae, or thick-headed flies, are an interesting and understudied group
of flies with a nearly worldwide distribution. They are only absent from
Antarctica and the islands of Pacific. The adults are nectarivorous and
the larvae are obligatory parasitoids of other insects. About 800 extant species
of conopids in 58 genera are known, and one fossil genus with two species has
been
recorded from Baltic amber (Stuke, 2005; Gibson and Skevington, 2013).
Stylogaster Macquart, 1835 is a peculiar genus of Conopidae, with 118 valid species
described so far, distributed amongst all biogeographic regions except for
the Palaearctic (Papavero, 1971; Camras and Parrillo, 1985; Smith and
Peterson, 1987; Yang, 1995; Camras and Parrillo, 1996; Stuke, 2006; Rocha
and Mello-Patiu, 2009; Schneider, 2010; Rocha and Mello-Patiu, 2012; Stuke,
2012; Pape and Thompson, 2013, Burt et al., 2014). The genus (thus the
subfamily) can be diagnosed by the following characters: a pronounced facial
keel, elongated prementum and labella, medial ommatidia larger than all
other conopid genera, females with an elongated tube-like terminalia, formed
by the segments VI, VII and VIII and dart-like eggs with a hard and scaled
chorion and barb-shaped spines (Kotrba, 1997). Stylogaster are parasitoids like all
other Conopidae but unlike the others, which are mostly parasitoids of
Hymenoptera, Stylogaster's preferred hosts are cockroaches (Blattaria) and grasshoppers
and their relatives (orthopteroids). Records of oviposition on calyptrate
flies have been described from Africa, but no evidence of development in
dipteran hosts has been found (Rettenmeyer, 1961; Couri and Pont, 2006;
Couri and Barros, 2010, Couri et al., 2013). In the New World and Africa,
many species are associated with army and driver ants, (Hymenoptera:
Formicidae: Ecitoninae), with Stylogaster attacking in the insects that run from the
swarms, but not the ants themselves (Lopes, 1937; Smith and Peterson, 1987; Skevington
et al., 2010).
Recent phylogenetic analyses give many alternative phylogenetic hypotheses
for Stylogaster: one as the sister group of all other conopids, or as sister of the
Lauxaniidae, depending on the method of analysis (Gibson et al., 2012;
Gibson and Skevington, 2013) or even a paraphyletic Conopidae, inside the
Sciomyzoidea, with Stylogaster separated from the other conopids (Wiegmann et al.,
2011). The age of Stylogaster has not been proposed before, as the only known fossil
conopid genus, Palaeomyopa Meulnier, 1912 is not closely related to Stylogaster (Hennig, 1966;
Gibson and Skevington, 2013). The short mouthparts with palps and widely
open cell R4+5 are putative plesiomorphic characters of Palaeomyopa, while the
presence of facial foveae and pad-like extensions of sternite 5 of the
female are shared with the non-stylogastrine conopids. Evolutionary studies
of Conopidae are important because this family is hypothesized to be the
sister taxon to all other schizophoran Diptera (Gibson et al., 2010), thus
providing insights into the evolution of diet and parasitoidism amongst the
enormous radiation of higher flies. The morphology of the mouthparts and
female terminalia suggests that ancestral conopids were nectarivorous and
parasitoids (Hennig, 1966), and must have evolved after the appearance of
the flowering plants. It remains unclear which hosts are plesiomorphic:
orthopteroids or hymenopterans, or another group entirely, unknown to date. The
estimated divergence from molecular data between schizophoran and
non-schizophoran Diptera is Upper Cretaceous (Santonian age), ca. 84.5 Mya
(Bertone and Wiegmann, 2009), although there were no conopids in their
analysis. A posterior analysis (Wiegmann et al., 2011), including conopids,
found that the Schizophora date from the K–T boundary, ca. 66 Mya. The oldest
schizophoran fossil is a Cretaceous (ca. 70 Mya, Maastrichtian age) puparium
from Alberta, Canada, Cretophormia fowleri McAlpine, 1970, and its family relationships are
uncertain. Except for this, all other fossil Schizophora are Cenozoic (ca. 66 Mya)
and very common in Baltic amber, although occurring in younger amber
(Grimaldi and Cumming, 1999; Grimaldi and Engel, 2005).
The early–middle Miocene amber-bearing sediments in the Dominican Republic are
associated with lignitic material and were formed in a great variety of
depositional environments, from coastal to deep marine. The age of the
amber-bearing sediments is estimated to be from Oligo-Miocene boundary (23 Mya)
to middle Miocene (14 Mya) (Iturralde-Vinent, 2001). The exact
formation of origin for the fossil is not provided on the specimen label,
but its age corresponds to the amber-bearing sediments cited by
Iturralde-Vinent (2001). This author suggests that those sediments were
likely formed from debris produced from active uplift of a highland
terrestrial or near-shore area.
This paper aims to describe the first known Stylogaster fossils, compare them with the
other conopid fossils, and discuss the systematic and biogeographic
relevance of this new species.
Material and methods
The examined specimens are from the American Museum of Natural History (AMNH)
collection.
They were observed with a Leica M80 stereomicroscope and then described.
Photographs were taken using a Canon EOS 50D camera with a Canon MP-E 65 mm
f/2.8 1–5x Macro Lens and a Canon Speedlite 430EX II flash. The camera was
attached to a StackShot (Cognisys Inc.) computerized rail, which was used to
take a series of images of the specimen. These images were then montaged
with Zerene Stacker version 1.04 (Zerene Systems LLC) to produce one image.
The montaged images were edited with Adobe Photoshop. The dorsum was
illustrated by tilting the specimen under various lighting conditions to
assess, capture and clearly display all the characters necessary for
diagnosis. The terminology used here follows McAlpine (1981) and Cumming and
Wood (2009), except when contradicted by Gibson et al. (2012), who are then
followed.
Results
Stylogaster grimaldii sp. nov.(Figures 1–6)Holotype: Dominican Republic amber, Oligo-Miocene, AMNH specimen
no. DR-15-411, 1 male; Paratype: Dominican Republic amber, Oligo-Miocene,
AMNH specimen (No DR no. given), 1.5 females.Type locality: Dominican Republic.Type stratum: Unknown.Etymology: Named after Dr. David Grimaldi, who has built an
impressive collection of amber for study at the American Museum of Natural
History. He obtained the specimens and brought them to our attention.
(a) Photo of male Stylogaster grimaldii sp. n., dorsal view. OT, ocellar
triangle. (b) Photo of male Stylogaster grimaldii sp. n., frontal view. Scale bar = 1 mm.
(a) Photo of male Stylogaster grimaldii sp. n., left lateral view. (b) Photo
of male Stylogaster grimaldii sp. n., right lateral view. MbT, mesobasitarsus. Scale
bar = 1 mm.
Male
Diagnosis: Pedicel and postpedicel subequal in size;
ocellar triangle short and pointed; mid basitarsus more than twice as long
as the remaining tarsomeres; hind trochanter and femur with long black setae
ventrally; tergite 1 with white setae laterally; tergite 5 longer than the
others; sternite 5 sclerotized.Description: Head: Frons yellow, fronto-orbital plate as wide as the
scape (Figs. 1a, 2a); gena and facial carina yellow, seeming to be covered by
microtomentum; prementum black at the middle and yellow at tips, basal
sheath of prementum clearly visible; ocellar triangle brown and rounded, not
reaching middle of the frons, ocellar tubercle prominent (Figs. 1a, 2a);
antennae entirely yellow, some setulae dorsally on anterior margin of scape;
pedicel and postpedicel subequal in size; arista brown, its third segment
enlarged at the base; head setae, per side, are 2 frontal, 6 orbital, 1
ocellar, 1 vertical and 2 occipital, all black (Fig. 2b); occiput and gena
covered by white setulae; ommatidia on medial side of the compound eyes
slightly enlarged (Fig. 1b).Thorax: Postpronotal lobe yellow, scutum light brown,
supraalar and posterior regions lighter; katepimeron and katepisternum of
the same colour as scutum, other pleural sclerites yellow; notopleuron with
strongly demarked suture, scutellum pointing upwards; proepisternal setae
white, all other black; setae on each side are 1 proepisternal, 1
postpronotal, 2 notopleural, 2 anepimeral, 2 supraalar, 2 dorsocentral, 2
postalar and 1 scutellar (Fig. 2a–b); fore and mid legs yellow; fore and mid coxae with long white setae apically, mid femur with a row of sparse black
setae ventrally; fore and mid tibiae coated with white setulae, which are
longer apically; fore basitarsus coated with white setulae; mid basitarsus
more than 2-fold longer than the other tarsomeres (Fig. 5); hind coxae light
brown, with long black setae apically, seeming to be covered with
microtomentum; hind trochanters with black setulae ventrally; hind femur
light brown in the tips and yellow in the middle, with moderately long black
setae ventrally at the base (Fig. 5); hind tibiae brown with white preapical
markings, and dense setulae at the apex; halter rounded, with the lateral
half of the knob brown and the medial yellow; wing with one basal seta,
costal setulae small; basal radial cell curved apically, M1+2 vein slightly
curved, forming a petiole with R4+5 (Fig. 2b).Abdomen: Abdominal tergites
yellow with light brown posterodorsal markings (Fig. 2b); tergites 2, 3 and
4 subequal in size, tergite 5 slightly larger than the others; tergite 1
with only white setae on the sides, covering all the lateral, tergite 2 with
a transversal series of 6 setae, 2 black and 4 white; white setulae present
in the anterior ventral margin of tergites 3, 4 and 5; long black setae in
the ventral margins of all tergites except tergite 1; sternites totally
membranous, but with visible limits.Terminalia: Mainly yellow; sternite 5 sclerotized,
with 2 transversal sets of black setulae; syntergosternite 7+8 nearly the
same size of the epandrium; cerci smaller than posterior surstyli and
apparently without hooked apex; cerci and posterior surstyli with black and
white setae (Fig. 2b).
Photo of 1.5 female Stylogaster grimaldii sp. n. right lateral view. Partial
specimen 1 is intact except for the tip of terminalia, while all that
remains of Partial specimen 2 is the abdomen with the terminalia intact.
Part. 1, 2. Partial specimen 1, 2. Scale bar = 1 mm.
Female
Description: Same as male, except for modified abdomen and
terminalia.Abdomen: Abdomen twice as long as thorax and head combined. Abdominal
tergites brownish-yellow, without clear maculation or markings (Figs. 3, 4);
tergites 2, 3 and 4 about equal in size, while tergite 5 slightly larger and
longer than others; tergite 6 somewhat constricted posteriorly and shorter
than 5, and appearing fused with tergite 7 (Figs. 3, 4, 6a); tergite 1 with
only white setae on the sides, tergite 2 with a transversal series of 4 long
black setae; white setulae absent from the anterior ventral margin of
tergites 3, 4 and 6; moderately long (shorter than in male) black setae
present in the ventral margins of all tergites except tergite 1; sternites
1–6 totally membranous, but with visible limits.Terminalia: Mainly dark brown; tergite
7 greatly elongated, cylindrical, slender, tube-like; tergite 8 virtually
indistinguishable, but appears fused to 7; tergite 9 indistinguishable but
also appears fused; tergites 7–8 with short, regular black setae dorsally,
ventrally and laterally; sternites appear fused; lateral lobes appear to
protrude from tergite 8, are short with rounded tips, cercus short with long
bushy black setae; hypoproct long (about 10 times as long as length of cercus),
and slightly rounded at tip, but arrow-shaped overall, covered in long black
setae, bushy (Fig. 6a–b).Comments: The other two known fossils of Conopidae have been
discussed by Hennig (1966), Camras (1994), Stuke (2005) and Gibson and
Skevington (2013); both are from Baltic amber. They are not associated
directly with extant taxa, but resemble the non-stylogastrine conopids in
having mobile female terminalia, pad-like appendices on the sternites and no
facial carina protruding. Stylogaster grimaldii is clearly a Stylogaster, having all the characters of the
group.Discussion: The Stylogaster fauna was known only from extant species until now. The discovery of a
fossil species from Oligo-Miocene sheds light regarding some questions about the
evolutionary history and biogeography of this group.
Photo of 1.5 female Stylogaster grimaldii sp. n. left lateral view. Partial
specimen 1 is intact except for the tip of terminalia, while all the remains
of Partial specimen 2 is the abdomen with the terminalia intact. Part.
1, 2. Partial specimen 1, 2. Scale bar = 1 mm.
Recent works on the phylogeny of Conopidae (Gibson et al., 2012; Gibson and
Skevington, 2013) place Stylogaster as the sister taxon of the remaining Conopidae. The
other fossil conopid group, Palaeomyopa, seems to be more related to the
non-stylogastrine conopids than to Stylogaster, although it has not been included in
any phylogenetic analysis. Palaeomyopa is known from Baltic amber dated from the early
Eocene, 44.1 to 47 Mya (Wolfe et al., 2009), suggesting that the separation
between Stylogaster and the Conopidae sensu Hennig (1966) has this minimum age. Stylogaster grimaldii probably
dates from the Oligo-Miocene boundary, ca. 23.03 Mya (International Commission
on Stratigraphy, 2013; Iturralde-Vinent, 2001).
Illustration of male Stylogaster grimaldii sp. n., dorsal view. Scale bar = 1 mm.
(a) Amalgamated illustration of female abdomen and
terminalia of Stylogaster grimaldii sp. n., right lateral view. Scale bar = 1 mm.
(b) Illustration of female terminalia of Stylogaster grimaldii sp. n., Partial specimen 2, right
lateral, slightly oblique view. Scale bar = 0.5 mm.
Stylogaster grimaldii was included in an all-species phylogeny
of Stylogaster, which divides the genus into two
monophyletic groups, one New World clade and one Old World clade (Rocha et
al., unpublished data). The New World group has a short ocellar triangle
clade and a long ocellar triangle clade. S. grimaldii morphologically resembles the
species with a short ocellar triangle, sharing with them many other
characters, such as the subequal-sized pedicel and postpedicel and the
absence of thick bristles on the epandrium. The hypothesized position of
other Stylogaster in this unpublished work is congruent with the molecular and
morphological cladogram of Gibson et al. (2012). The hypothesized position
of the fossil species, as the sister of all other species with a short
ocellar triangle, indicates that the separation between Old/New World and
long/short ocellar triangle New World clades is probably older than the
Neogene (23.03 Mya). The position of S. grimaldii adds little to resolve the conflict
concerning the placement of Conopidae within Schizophora (Gibson et al.,
2010). Different analytical methods used by Gibson et al. (2010) produced
different hypotheses, with Parsimony suggesting that conopids are sister to
Schizophora and Bayesian suggesting that Conopidae are sister to Lauxaniidae
Macquart. Fossil Lauxaniidae are rare (Evenhuis, 1994) but have been found
in Baltic amber (Evenhuis, 1994; Poinar Jr., 1992) and in the Fushun amber of
China (Szwedo et al., 2013; Hong, 1981). The Fushun amber deposit has
been dated to the Paleogene, specifically the Paleocene and early Eocene
epochs (50–53 Mya) (Wang et al., 2014). Finding an older conopid fossil (in
the 70 Mya range) would add considerable support to the conopid–Schizophora
sister hypothesis. Given that the minimum age of conopids (47.9 Mya based on
Palaeomyopa) is similar to that of the oldest lauxaniid, both hypotheses remain viable.
Another fossil Stylogaster in Dominican amber, a female, is owned by a private
collector in Italy (D. Grimaldi, personal communication, 2014) but could not be obtained at this
time. A photo of the specimen can be seen at
http://www.terratreasures.com/amber/flagship/dr5705stylogaster/DR5705stylogaster.htm,
but its relationship to S. grimaldii is not entirely clear, because the ocellar
triangle is not visible. However, the short postpedicel and long female
terminalia suggest it is related to S. grimaldii. If the identity of this
Dominican fossil is confirmed as another new species, then this suggests that there was greater
diversity in the Antilles because the only extant species recorded from these
islands is S. iviei Camras, 2003. The fact that at least two lineages are
represented by these taxa suggests that the West Indies was an important
corridor and has lost much of its fauna through extinction events (Rocha et
al., unpublished data). The insights coming from this fossil species
illustrate the contribution that fossil taxa can make with respect to
conopid phylogenetics.
L. S. G. Rocha analysed the specimens and wrote the descriptions and the main text; T. O. Burt
revised the text, especially the conclusions, and made the drawings of the
specimens; C. A. de Mello-Patiu and J. H. Skevington revised the text and made major corrections to the
primary manuscript.
Acknowledgements
Funding for L. S. G. Rocha came from the Sandwich fellowship programme of CAPES
(Coordenação de Aperfeiçoamento de Pessoal de Nível
Superior, Brazil, proc. no. 10454-12-8) and FAPERJ – Fundação Carlos
Chagas Filho de Amparo à Pesquisa do Estado do Rio de Janeiro, Brazil
(proc. no. E-26/110.409/2012). Agriculture and Agri-Food
Canada supported L. S. G. Rocha during his work in Ottawa and provide operational
funds to J. H. Skevington. We thank David Grimaldi for providing the specimen to us,
Jens-Hermann Stuke and Jeff Cumming for guidance and Michelle Locke for
taking the stereomicroscopic images of the male specimen. This work is part
of the PhD thesis of L. S. G. Rocha.
Edited by: D. Korn
Reviewed by: J.-H. Stuke and S. Marshall
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