Brachymeria mochica, a new Neotropical species of Chalcididae (Hymenoptera: Chalcidoidea) discovered on the archaeological site of Huacas de Moche, Peru with a review of related species
Author
Delvare, Gérard
Author
Huchet, Jean-Bernard
text
Zootaxa
2017
4290
1
43
60
journal article
32723
10.11646/zootaxa.4290.1.2
a2bf715a-07c1-4693-adbb-f8f85b8d8faa
1175-5326
828897
8172109F-E545-4EB7-9E9D-D91E7DD1FA42
Brachymeria podagrica
(Fabricius, 1787)
Chalcis podagrica
FabriciUS 1787: 272. Original deScription ♂. India, TranqUebar. Lectotype by
BoUček 1972
: 241
.
Brachymeria podagrica
(FabriciUS):
BoUček 1972
: 241
.
=
Chalcis femorata
NeeS 1834: 412. Original deScription. Germany. Synonymy by
Dalla Torre 1898
: 391
. Type apparently deStroyed.
=
Chalcis Fonscolombei
DUfoUr 1841
: 16
–
17. Original deScription ♀. SoUthern France. Lectotype by BUrkS (1936). Synonymy by
BoUček 1972
: 241
.
=
Chalcis alphius
Walker 1846a
: 108
. Original deScription ♂. India, Bombay. Lectotype and Synonymy by
BoUček 1972
: 241
.
=
Chalcis xerxena
Walker 1846a
: 108
–
109. Original deScription ♂. India, Bombay. Lectotype and Synonymy by
BoUček 1972
: 241
.
=
Chalcis restituta
Walker 1862
: 351
–
352. Original deScription Jamaica. Lectotype and Synonymy by
BoUček & DelVare 1992
: 30
.
=
Chalcis pulchripes
Holmgren 1868
: 436
–
437. Original deScription ♂. PhilippineS, Manila. Lectotype and Synonymy by
BoUček 1972
: 241
.
=
Chalcis mansueta
Walker 1871
: 48
–
49. Original deScription ♀. Hong-Kong. Lectotype and Synonymy by
BoUček 1972
: 241
.
=
Chalcis callipus
Kirby 1883
: 75
. Original deScription ♀. Japan. Lectotype and Synonymy by
BoUček 1972
: 241
.
=
Chalcis mikado
Cameron 1888
: 117
. Original deScription ♀. Japan. Lectotype and Synonymy par
BoUček 1972
: 241
.
=
Chalcis eccentrica
Cameron 1897
: 39
. Original deScription ♂. India, Bombay. Lectotype and Synonymy by
BoUček 1972
: 241
.
=
Chalcis ferox
Kieffer 1905
: 263
–
264. Original deScription ♀ ♂. ReUnion ISland [BoUrbon] and MadagaScar, NoSSi-Bé.
Syn. nov.
Synonymy With
B.
fonscolombei
by
Steffan 1959
: 42
.
=
Chalcis ferox
Var.
coxalis
Kieffer 1905
: 264
. Original deScription ♀. MadagaScar, NoSSi-Bé.
Syn. nov.
Synonymy With
B.
fonscolombei
by
Steffan 1959
: 42
.
=
Chalcis borneanus
Cameron 1905a
: Original deScription ' ♀'. Borneo. Lectotype ♂ and Synonymy by
BoUček 1972
: 241
.
=
Chalcis spilopus
Cameron 1905b
: 231
. Original deScription ♀. SoUth
Africa
, TranSVaal.
Syn. nov
.
=
Chalcis capensis
Cameron 1905b
: 311
–
312. Original deScription ♀. SoUth
Africa
, Katberg.
Syn. nov
.
=
Chalcis transvaalensis
Cameron 1911
: 214
–
215. Original deScription ♀. SoUth
Africa
.
Syn. nov
.
=
Tumidicoxoides kurandaensis
GiraUlt 1913
: 86
. Original deScription ♀. AUStralia, QUeenSland. Synonymy by
BoUček 1988
: 72
.
=
Chalcis dipterophaga
GiraUlt & Dodd in
GiraUlt 1915
: 320
–
321. Original deScription ♀. AUStralia, QUeenSland. Synonymy by
BoUček 1988
: 72
.
=
Tumidicoxoides paucipunctatus
GiraUlt 1915
: 326
. Original deScription ♀. AUStralia, Northern Territory. Synonymy by
BoUček 1988
: 72
.
=
Chalcis neglecta
MaSi 1916
: 84
. Original deScription ♀. Italia, Giglio.
Syn. nov
. Synonymy With
B.
fonscolombei
by
Steffan 1959
: 42
.
=
Chalcis sodalis
MaSi 1917
: 128
–
129. Original deScription ♀ ♂. SeychelleS ISlandS: Mahé, SilhoUette.
Syn. nov
. Synonymy With
B.
fonscolombei
by
Steffan 1959
: 42
.
=
Chalcis vegai
GiraUlt 1924
: 175
. Original deScription ♀. AUStralia, QUeenSland. Synonymy by
BoUček, 1988
: 72
.
=
Chalcis garutianus
Günther in
Hellen &Günther 1936
: 73
–
74. Original deScription ♀. JaVa, Kamodjan. Lectotype and Synonymy by
BoUček 1972
: 241
.
=
Brachymeria Fonscolombei
Var.
gananensis
MaSi 1938
: 210
–
212. Original deScription ♀. Somalia, Belet-Amin and MogadiScio.
Syn. nov
.
=
Chalcis vulcani
Schmitz 1946
: 63
–
64. Original deScription ♂. RUanda, BUrambi (Volcan MUhaVUra).
Syn. nov
.
=
Brachymeria aligarhensis
Husain & Agarwal 1982
: 499
–
501. Original description ♀. India, Uttar-Pradesh. Synonymy by
Narendran 1989
: 260
.
Material examinated.
Type material
.
Chalcis Fonscolombei
.
Lectotype, a pinned ♀ from the Dufour collection, identified by Steffan (MNHN) (
Figs 29, 30
).
Chalcis capensis
. Lectotype, a ♀ on a minutien pin, the designation of which by Bouček (1971) is validated here (
Fig. 33
) (BMNH).
Chalcis transvaalensis
. Lectotype, a ♀ in several pieces, remounted on a rectangular card, the designation of which by Bouček (1971) is validated here (
Fig. 34
) (BMNH).
Chalcis spilopus
. Lectotype, a ♂ on a minutien pin, the designation of which by Bouček (1971) is validated here (BMNH, Hym. type 5.155).
Chalcis ferox
. Lectotype, a ♂ on a minutien pin, the designation of which by Rasplus is validated here; it is labelled 'Bourbon [manuscript] /
Chalcis ferox Sauss.
type ♂' [manuscript, in Kieffer's handwriting] (
Fig. 35
) (MNHN). The ♀ quoted by
Kieffer (1905)
from Nossi Bé could not be retrieved.
Chalcis ferox
var.
coxalis
.
Lectotype, a pinned ♂ labelled 'Madag.' [manuscript] / '
Ch. ferox Sss.
' [manuscript] / '
Chalcis ferox Ss.
var.' [manuscript, in Kieffer's handwriting] (
Fig. 36
) (MNHN).
Chalcis neglecta
. Lectotype, a ♀ glued on a rectangular card, the designation of which by Bouček (1971) is validated here; it is labelled 'Is. Giglio vii 901 G. Doria' / '
Chalcis neglecta
Masi 1916
typus!' [manuscript, in Masi's handwriting] / ' ♂
Brachymeria neglecta (Masi)
=
podagrica
(F.) Bouček det. 1971' [manuscript, in Bouček's handwriting] (
Fig. 37
) (MCSN).
Brachymeria Fonscolombei
var.
gananensis
. Lectotype, the designation of which by Bouček (1971) is validated here, labelled 'EL DUGUL Somalia Italiana E. Confalonieri' / '
Brachymeria Fonscolombei
subsp. gananensis Masi ♀ det. L. Masi' [manuscript in Masi's handwriting] / 'co-typus' / 'Lectotype' [in BMNH format] with 'Bčk 1971' in the underside (
Fig. 38
) (MCSN).
Chalcis vulcani
. Holotype ♂, by original designation (MRAC, n° 567).
Other material. PERU.
Trujillo
,
Huaca de la Luna
,
08°08′06″S
78°59′29″W
,
56 m
, 06.v/
01.vi.2009
,
in fly trap
, J.-
B. Huchet
and
A. Chauchet
leg. (
1 ♀
GDPC
)
.
FRANCE
.
Bouches-du-Rhône
,
Istres
,
01.vii.2003
,
P. Ponel
leg. (
1 ♂
GDPC
)
;
Gard
, Le-Grau-du-Roi,
l'Espiguette
beach,
24.v.1997
,
G. Delvare
leg. (
1 ♀
GDPC
)
;
Hérault
,
Agde
,
Rochelongue
,
29.vii.1979
, J.-
M. Maldès
leg. (
1 ♀
GDPC
)
;
Montpellier
, campus
ENSA
, on mackerel corpse,
15.ix.2000
,
M. Martinez
leg. (
3 ♀
GDPC
)
;
same locality and collector, ex
Sarcophaga proxima
Rondani
[
Sarcophagidae
] on a snake carcass, adult emergence 10/
13.vi.2003
(
2 ♀
,
11 ♂
GDPC
)
;
Lot-et-Garonne
,
Buzetsur-Baïsse
,
30.v.1995
,
Malaise trap
, J.-
P. Sarthou
leg. (
1 ♀
GDPC
)
.
SPAIN
.
Tarragona
,
Cambrils-de-Mar
,
30.v.1990
,
H. Tussac
leg. (
2 ♂
GDPC
)
.
Comment.
Steffan (1959)
previously synonymised
C. ferox
Kieffer
and its variety
C. ferox coxalis
with
B.
fonscolombei
, as well as
C. neglecta
Masi.
Brachymeria fonscolombei
was itself synonymised later with
B.
podagrica
by
Bouček (1972)
. The name
podagrica
was known from
Dufour (1841)
but because the
type
was described from
India
he thought that the European specimens belonged to another species, which he named
Chalcis Fonscolombei
in honour of Boyer de Fonscolombe. This name was adopted by subsequent authors until
Bouček (1972)
synonymised it with
B.
podagrica
.
Brachymeria
.
podagrica
was sometimes mixed with
B. parvula
(Walker) under its junior synonym
Chalcis Dalmanni
Thomson
, especially by Stefani (1889) because the females of both species have the metafemur mostly reddish. It is nevertheless evident that Stefani dealt with
B.
podagrica
because he reared his specimens from flies emerging from carrion while
B. parvula
is a secondary parasitoid of locusts.
Diagnosis.
Apex of scape reaching base of median ocellus. Antennal flagellum mostly linear, hardly fusiform or clavate and bearing short erect setae only (
Fig. 21
). Mesonotum with moderately dense puncturation, the interspaces between punctures narrower that the points themselves (
Fig. 22
). Propodeum without spiracular callosity. Metacoxa and metafemur reddish, the latter with a small ivory spot apically. Metafemur slender 1.85
–
1.95× as long as broad, with an acute inner tooth basally; its outer surface with relatively dense puncturation, the basal piliferous points hardly more distant from each other than their own diameter (
Fig. 24
). Metatibia reddish, respectively with a subbasal and dorsoapical ivory spots. Gaster acuminate, 1.8
–
2.0× as long as broad (
Fig. 23
).
Hosts and biology.
Table 1
lists the known hosts of the species and the authorships of the observations. Dubious host records are not included in the table, especially those for the hyperparasitoids of locusts which, as already mentioned by
Steffan (1959)
, result from past confusion with
B. parvula
. We also omitted the hosts quoted for
B.
amenocles
, which until now was wrongly mixed with
B.
podagrica
following
Bouček (1972)
(see below).
FIGURES 21–28. 21‒24,
Brachymeria podagric
a, ♀.
21
, antenna exclUding Scape.
22
, meSoSoma in dorSal VieW.
23
, metaSoma in dorSal VieW.
24
, metafemUr.
25‒28,
Brachymeria amenocles
, ♀.
25
, antenna exclUding Scape.
26
, meSoSoma in dorSal VieW.
27
, metaSoma in dorSal VieW.
28
, metafemUr.
FIGURES 29–39. 29
, type SerieS of
Brachymeria Fonscolombei
DUfoUr (MNHN).
30–39
, labelS of primary typeS.
30
,
Brachymeria Fonscolombei
.
31
,
Chalcis varipes
Walker.
32
,
Chalcis amenocles
Walker.
33
,
Chalcis capensis
Cameron.
34
,
Chalcis transvaalensis
Cameron.
35
,
Chalcis ferox
Kieffer.
36
,
Chalcis ferox
Var. Kieffer.
37
,
Chalcis neglecta
MaSi.
38,
Brachymeria Fonscolombei
Var.
gananensis
MaSi.
39
,
Brachymeria Beccarii
MaSi.
Sarcophagidae
(18 species) are predominant in the list of known hosts, which also includes a number of
Calliphoridae
(11 species) together with a few
Muscidae
. Cultures made by several authors (Stefani 1889;
Parker 1924
;
Roberts 1933
;
Carlos 2014
) showed that females emerging from sarcophagid hosts were always much more numerous compared to those that developed at the expense of the two other dipterous families. These flies often feed on corpses of various vertebrates, primarily mammals, but also from snakes (Beaver 1936; Michel Martinez pers. com.) and even fishes, as shown by the trapping of
B.
podagrica
by exposing mackerel corpses (Michel Martinez, pers. com.). Some blowflies, such as
Cochliomyia macellaria
(Fabricius) (Calliphoridae)
, also figure in the spectrum of hosts. The presence of individuals along the Mediterranean seaside (see above, examined material) suggests that
B.
podagrica
is normally resident in this habitat, possibly searching for larval stages of flies developing at the expense of fish corpses washed up on the shore.
TABLE 1.
Known hosts of
Brachymeria podagrica
. With a very few exceptions, only data from original publications are included, hence excluding second-hand citations in catalogues or data.
HoSt SpecieS HoSt family Reference and diStribUtion
Ophyra aenescens
(Wiedemann, 1830)
MUScidae
Marchiori
et al
. 2003a
and
2003c
:
Brazil
Ophyra
Sp.
MUScidae
CarloS 2014
:
Brazil
Synthesiomyia nudiseta
(WUlp, 1883)
MUScidae
RobertS 1933
:
USA
Philornis
Sp.
MUScidae
CoUri
et al
. 2006
:
Brazil
Calliphora coloradensis
HoUgh, 1899
Calliphoridae
RobertS 1933
: USA
Calliphora grahami
Aldrich, 1930
Calliphoridae
HabU 1960: Japan
Cochliomyia macellaria
(FabriciUS, 1775)
Calliphoridae
RobertS 1933
:
USA
Chrysomya albiceps
(Wiedemann, 1819)
Calliphoridae
Marchiori
et al
. 2002b
and
2003a
;
CarloS 2014
:
Brazil
Chrysomya megacephala
(FabriciUS, 1794)
Calliphoridae
CarloS 2014
:
Brazil
Hemilucilia flavifacies
Enderlein, 1931
Calliphoridae
Marchiori
et al
. 2003b
;
CarloS 2014
:
Brazil
Lucilia caesar
(LinnaeUS, 1758)
Calliphoridae
BUrkS 1960:
USA
Lucilia illustris
(Meigen, 1826)
Calliphoridae
BUrkS 1960:
USA
Phaenicia mexicana
(MacqUart, 1843)
Calliphoridae
RobertS 1933
:
USA
Phaeanicia sericata
(Meigen, 1826)
Calliphoridae
RobertS 1933
:
USA
:
OliVa 2008
:
Argentina
Phormia regina
(Meigen, 1826)
Calliphoridae
BUrkS 1960:
USA
Kellymyia plinthopyga
(Wiedemann, 1830)
Sarcophagidae
RobertS 1933
:
USA
Kellymyia impar
(Aldrich, 1916)
Sarcophagidae
RobertS 1933
:
USA
Kellymyia plinthopyga
(Wiedemann, 1830)
Sarcophagidae
RobertS 1933
:
USA
Oxysarcodexia thornax
(Walker, 1849)
Sarcophagidae
Marchiori
et al
. 2002a
and
2003a
; GonçalVeS
et al
. 2012;
CarloS 2014
: Brazil
Peckia chrysostoma
(Wiedemann, 1830)
Sarcophagidae
Marchiori 2001
;
Marchiori
et al
. 2003a
;
CarloS 2014
:
Brazil
Sarcophaga africa
(Wiedemann, 1824)
Sarcophagidae
DUfoUr 1841
:
France
[Under
S. haemorrhoidalis
]:
RobertS
1933
:
USA
Sarcophaga argyrostoma
(RobineaU-DeSVoidy, 1830)
Sarcophagidae
UrSU & TUdor 1975; AndrieScU 1988:
Romania
;
OliVa 2008
:
Argentina
Sarcophaga carnaria
(LinnaeUS, 1758)
Sarcophagidae Stefani 1889
: Italy: Sicily; BUrkS 1960: USA
Sarcophaga crassipalpis
MacqUart, 1839
Sarcophagidae
OliVa 2008
: Argentina
Sarcophaga dux
ThompSon, 1869
Sarcophagidae
ChoWdhUry & HoWlader 1975
:
BangladeSh
Sarcophaga exuberans
Pandellé, 1896
Sarcophagidae
ChoWdhUry & HoWlader
1975
and
1978
:
BangladeSh
Sarcodexia lambens
(Wiedemann, 1830)
Sarcophagidae
Marchiori
et al.
2003a
; GonçalVeS
et al
. 2012;
CarloS 2014
:
Brazil
Sarcophaga musitasi
CUrran, 1934
Sarcophagidae
Fry 1989
: PakiStan
Sarcophaga peregrina
RobineaU-DeSVoidy, 1830
Sarcophagidae
HabU 1960: Japan
Sarcophaga proxima
Rondani, 1860
Sarcophagidae Martinez
perS. comm.:
France
(preSent pUblication)
Sarcophaga
Sp.
Sarcophagidae
Parker 1924
:
France
(cUltUre)
Sarcophaga
Sp.
ex corpSe
Achatina fulica
Sarcophagidae
BeaVerS 1986:
Thailand
Sarcophaga tuberosa
(Pandellé, 1896)
Sarcophagidae
FabritiUS & AndrieScU 1987; AndrieScU 1988:
Romania
As is the case for the other species belonging to the
minuta
group,
B.
podagrica
is a solitary larval-pupal endoparasitoid (
Dufour 1841
; Stefani 1889;
Parker 1924
;
Roberts 1933
). The latter author studied in detail the biology, ethology and phenology of
B.
podagrica
, collecting copious samples and conducting large scale rearing of flies with the aim of evaluating the potential of the chalcidid to regulate the populations of the blowfly. All the above authors found a clear preference of
B.
podagrica
for the pre-imaginal stages of
Sarcophagidae
relative to those of
Calliphoridae
, often rejecting the latter family. This might be explained by the fact that the adults of the
Brachymeria
, having emerged from the puparia of
Calliphoridae
, are much smaller in size—sometimes only half the size—than those emerging from
Sarcophagidae
, apparently relating to differences in the size of the hosts themselves (
Roberts 1933
).
Females of
B.
podagrica
oviposit within the body of the last larval stage of the host, which survives until pupation. Oviposition is especially induced by the presence of fluids issuing from the animals attacked by the flies, whether from living tissues or decaying corpses. The female uses her posterior legs to hold the larva, which promptly reacts to her attack. Reproduction is of the preovigenic
type
; the single mating occurs very soon after emergence of the imagos and is quickly followed by oviposition. This might be an adaptation to the presence of a large number of host larvae, grouped into clusters on decaying carrion. In temperate regions
B.
podagrica
diapauses during winter in the last larval or prepupal stage, within the host puparium. Several generations occur during summertime (
Roberts 1933
).
Distribution.
Brachymeria podagrica
is a cosmopolitan species, recorded from 45 countries (Noyes 2016), distributed from
Australia
to
Sweden
and
Argentina
. In the Neotropical Region the chalcidid is recorded from
Mexico
,
Haiti
,
Jamaica
,
Peru
,
Brazil
and
Argentina
. Its region of origin is not known and it is possible that is was introduced by Europeans when colonising the Americas.
Chalcis amenocles
Walker
and
Brachymeria Beccarii
Masi
were synonymised with
B.
podagrica
by
Bouček (1972)
. However, examination of their
types
show that they belong to another distinct and valid species which is revalidated here.