PEPSINAE
Pepsinae
is also a diverse group with a conflicting history of classification, and several genera of uncertain membership. For example,
Epipompilus
was previously considered a monotypic subfamily ( Shimizu, 1994), and then transferred to
Ctenocerinae ( Pitts et al., 2006)
. More recently, cladistic morphological analyses with qualitative and quantitative characters suggested
Epipompilus
to be the sister to
Minagenia Banks
(E. F. Santos, pers. comm.).
Minagenia
has suffered similar inconsistencies.
Minagenia species
are morphologically homogeneous, but difficult to assign to a subfamily ( Dreisbach, 1953). Townes (1957) placed
Minagenia
in
Ceropalinae
; Haupt (1959), Evans (1973), and Pitts et al. (2006) considered it a member of
Pepsinae
. Another example concerns the variable
Chirodamus Haliday. Roig Alsina (1989)
split
Chirodamus
into six Neotropical genera:
Chirodamus
s.s.,
Plagicurgus Roig Alsina
,
Calopompilus Ashmead
,
Pompilocalus Roig Alsina, Aimatocares Roig Alsina
, and
Anacyphononyx Banks.
Chirodamus
s.s. was placed in
Pompilinae
by Pitts et al. (2006), but the other genera of
Chirodamus
s.l. have been considered as
Pepsinae
.
Our results recovered a monophyletic
Pepsinae
in the relaxed-clock analysis only, with good support. Most of the deeper relationships within this clade were not supported, whereas several lineages of more recent origin were highly supported. The molecular phylogeny supports the assignment of the controversial genera, discussed above, as members of
Pepsinae
.
Epipompilus
is monophyletic, although its position within
Pepsinae
is ambiguous. It has a disjunct distribution, with species found in the Neotropics and Australasia. In our molecular phylogeny and in a morphological phylogenetic study (E. F. Santos, pers. comm.),
Epipompilus
is recovered as two major clades, one Neotropical and the other Australasian.
Epipompilus
hunt spiders inside their burrows and permanently paralyse them before oviposition ( Pollard, 1982).
Our analyses also support
Minagenia
and
Chirodamus
s.l. as members of
Pepsinae
.
Minagenia
is strongly supported as monophyletic, but its position within
Pepsinae
is uncertain. Species of
Minagenia
differ from other
Pepsinae
by having a straight stinger, a compressed metasoma, bifid claws and the veins 2 r-m and 3 r-m continuously curved outward and with similar appearance. They are ectoparasitoids, paralysing their prey only temporarily. Our results also confirm Roig Alsina’s (1989) division of
Chirodamus
into several genera (to the extent that we have sampled these taxa).
Amongst
Pepsinae
tribes, the most morphologically and behaviourally diverse is
Ageniellini
(clade L, excluding
Cyphononyx
). The monophyly of
Ageniellini
was recovered by Shimizu (1994), Pitts et al. (2006), and Shimizu, Wasbauer & Takami (2010), but this tribe is made paraphyletic in our analyses by the position of
Melanagenia
.
Melanagenia
was recently described by Wahis, Durand & Villemant (2009), and was defined and placed in
Ageniellini
by having the metasoma petiolate and by the first tergite lacking a transverse carina. Our results indicate that
Melanagenia
is unrelated to other
Ageniellini
. Rather, it emerges as sister to
Sphictostethus
, with which
Melanagenia
shares states of facial characters (lack of malar space with eyes touching mandibles and a clypeus somewhat rectangular and convex), pronotal characters (rounded with a deep sulcus laterally), and wing-venation characters. However, as
Melanagenia species
lack a carina on the first tergite and have a petiolate metasoma, these two character states – although useful in identifying
Ageniellini
taxa – can no longer be considered unique synapomorphies of the tribe. The observation that
Phanagenia Banks
(
Ageniellini
) possesses a carina on the first metasomal segment further undermines the diagnostic value of this metasomal character.
Melanagenia
is herein removed from
Ageniellini
and placed in
Pepsini
. As discussed above (see
Ctenocerinae
),
Lepidocnemis
is sister to
Pompilocalus
and Aimatocares, within a larger lineage including
Sphictostethus
and
Melanagenia
.
Lepidocnemis
is the only representative of Neotropical
Ctenocerinae
in our study and is herein transferred to
Pepsinae
.
Pepsini
and the other tribes are in dire need of further studies and redefinition of most of their taxa. Our samples and analyses are not sufficient to make further nomenclatural decisions regarding tribes.
Pepsinae
(clade D) are now defined by: (1) metasomal sternum 2 with a distinct sharp transverse groove; (2) the mesofemur and the metafemur without subapical spine-like setae set in grooves or pits; (3) the metatibia with apical spine-like setae of uniform length, the setae not splayed; and (4) the fore wing with vein Cu1 simple at base, without any definite downward deflection, such that the second discal cell (2D) is without a ‘pocket’ posterior. A broad range of nesting behaviour occurs within this subfamily, including nesting in preexisting cavities, using the spider’s burrow, digging a burrow in the ground, building nests of mud, and behaving as true parasitoids and kleptoparasites.
