Fauna of NZ 45 - Landcare Research

54 Kuschel (2003): Nemonychidae, Belidae, Brentidae (Insecta: Coleoptera)
consenus is shown in Fig. 191) Belini is monophyletic and
an SAW analysis resulting in 9 trees (by two iterations,
consensus tree shown in Fig. 192) resulted in a
monophyletic Belini which is sister group to the remaining
taxa which includes a monophyletic Pachyurini. Using
these exemplar taxa, we added the adult characters which
resulted in 1 tree (TL = 83, CI excluding uninformative
characters = 0.63, RI = 0.6824) shown in Fig. 193. This
tree shows a monophyletic Agnesiotidini which is sister
taxon to the remaining taxa which includes a monophyletic
Belini.
A completely combined analysis (all taxa and characters)
resulted in 536 trees (TL = 144, CI = 0.42, RI = 0.76) with
the same relationships shown in Fig. 188 (the SAW analysis
resulted in the same trees as shown in the consensus tree in
Fig. 189). A priori weighting of characters 9, 23, 28, 30, 36,
43, and 46 resulted in 168 trees (TL = 209, CI = 0.59, RI =
0.89) that are similar to those seen with in Fig. 190 but
with poorer resolution (consenus tree in Fig. 194). A
consenus of 12 trees found by SAW is shown in Fig. 195.
Agnesiotidini
To this group we add Atractuchus from Pachyurini and
formally remove Agathobelus, Apagobelus, Basiliobelus,
and Macrobelus, which are transferred to Pachyurini. The
Agnesiotidini is monophyletic based on the following
characters (reference tree in Fig. 189 and unique characters
indicated by an *): shape of temples in dorsal view parallelsided
or broadening caudad (1-1, present also in Belini),
sides of frons near eyes with superficial or well defined
grooves (6-1), third antennomere reaching eyes (14-1 r ,
reverses in Atractuchus and Dicordylus marmoratus),
surface of scutellum convex (20-2*), base of elytra in profile
relative to pronotum higher (23-1*), shape of radial cell
window of hind wing pentagonal or tetragonal (28-1*),
extension of Cu in relation to hind margin of wing ending
some distance from it (29-1, present outside this group in
Trichophthalmus), sides of pro- and mesocoxae completely
closed (36-1*), crenulated ridge on laterodorsal surface of
tibiae present (38-1, present outside this group in all Belini),
one spine present on lower edge on one or more femora
(42-1, present outside this group in some Belini), length of
parameral sector relative to apodeme distinctly longer (46-
1*). The following characters are concordant with these,
including larval characters which are coded as missing for
most terminals: sides of prothorax at apex in dorsal view
strongly constricted to form a collar (18-1, equivocal
ancestral reconstruction), median suture absent (54-1), and
spiracles bicameral (55-1). If Agnesiotidini are a derived
group in Pachyurini (Fig. 190), the following characters
support its monophyly: apex of epistome truncate or
bilobed (8-0), ommatidia coarse (11-0 r , reverses in
Agnesiotis), (20-2*), (22-0*), (23-1*), (28-1*), (29-1),
crossvein a 2 -a 3 present as a relatively long vein (31-0,
DELTRAN), vein A 1 interrupted or absent (33-1), (36-
1*), (38-1), (42-1), and (46-1*).
Dicordylus marmoratus is more closely related to
Atractuchus annulifer than it is to D. binotatus, according
to characters 4-0, 12-0, 19-2, and 26-0, which would suggest
that these two species could be merged under one genus or
that an additional genus would have to be proposed for D.
marmoratus. We adhere to the synonymy of the two
genera as stated previously as their differences are too
slight to warrant independent status, although appearing
in the trees as separate taxa. Though Agathinus groups as
sister taxon to Cyrotyphus in one case it should not be
included in this genus (see Remarks in taxonomic section.
p. 28).
Belini
Belini is a monophyletic group based on (reference tree in
Fig. 190): shape of temples in dorsal view parallel-sided or
broadening caudad (1-1, present also in Agnesiotidini),
shared track of A 1 +A 2 apicad of a 2 -a 3 or junction of the
shared track with A 3 short (32-1*), position of procoxae
slightly antemedian (35-1*), length of hind femora in relation
to hind margin of ventrite 2 not or just reaching it (37-
1*), crenulated ridge on laterodorsal surface of tibiae present
(38-1, present outside this group in all Agnesiotidini),
aedeagus with pedon and tectum at least partially fused to
a tubular structure (43-1*), insertion of antennae postmedian
(13-1, equivocal ancestral reconstruction), and ventral
or subterminal placement of anus (56-1, larval character
that is coded as missing for most terminals) are concordant
with these. If Belini is derived within Pachyurini (Fig.
189), then the following characters support its monophyly:
(1-1), apical tooth of mandibles in dorsal view exposed in
front of median tooth (9-0*), elytral declivity at apex in
profile absent or gentle (24-1 r , reverses in Homalocerus,
Isacantha, and Trichophthalmus), (32-1*), (35-1*), (37-
1*), (38-1), (43-1*), mandible adentate (49-2, unique and
probably specialised in Rhinotia), tormae absent (50-1,
unique and probably specialised in Rhinotia), occipital
foramen closed (53-0, shared with Agnesiotidini).
Pachyurini
As stated above to this group we add Agathobelus,
Apagobelus, Basiliobelus, and Macrobelus from
Agnesiotidini, and formally transfer Atractuchus to
Agnesiotidini. By fixing Pachyurini as a monophyletic
group in MacClade, this group is supported by the absence
of a crenulated ridge on laterodorsal surface of tibiae