Rab. Leschen, Systematics of Nitidulinae (Coleoptera : Nitidulidae): phylogenetic relationships, convexity and the origin of phallalophagy, INVERTEBR T, 13(5), 1999, pp. 845-882
Nitiduline beetle genera of the Cyllodes complex are studied in the context
of a phylogenetic analysis of 19 ingroup and eight outgroup taxa and 63 ch
aracters derived from adult morphology and host relationship. In this analy
sis, the Cyllodes complex is shown to be monophyletic. By eliminating the p
roblematic taxon Camptodes and a priori weighting of the presence/absence o
f procoxal rests, two monophyletic taxa are obtained within the Cyllodes co
mplex (Cyllodes and Oxycnemus groups). A phylogenetic analysis of 13 termin
al taxa represented by larval and adult semaphoronts shows a monophyletic C
yllodes complex in one of two trees. The Cyllodes complex is formally recog
nised as Cyllodini Everts, containing two or more monophyletic groups that
are recognised on an informal basis. The Oxycnemus group is well supported
and may include the enigmatic genus Camptodes. Taxonomic actions include tw
o synonymies under Apsectochilus Reitter (= Meoncerus Sharp, syn. nov.; = L
ordyrops Reitter, syn. nov.) and type species designations for Neotropical
genera described by David Sharp (Apsectochilus, Eusphaerius, Mecyllodes, an
d Somatoxus). Carinocyllodes, gen. nov. (type species: Eusphaerius lubricus
Leschen & Carlton) is described and includes two additional species (C. be
lli, sp. nov. and C. depressus, sp. nov.).
Convexity is a character complex in nitidulids, and must be partitioned for
phylogenetic reconstruction and character analysis. Two classes of charact
er variation (dependent and independent) are discussed with regard to the e
volution of convexity in the context of cladograms. A highly convex form wi
th the capability to roll into a ball by tucking the head beneath the proth
orax has evolved at least one or more times in Nitidulidae and several time
s in unrelated beetle families.
Fungal host-use patterns are highly concordant with the phylogenetic branch
ing pattern in Cyllodini. Most members are associated with large-bodied Bas
idiomycetes, with the exception of Camptodes which occurs on flowers and de
velops in rotten vegetation. Most species of the Oxycnemus group are restri
cted to Phallales, an obligate host pattern that is old relative to those a
ssociations in the remaining Cyllodini that occur on other fungi. Another s
ignificant shift occurred from phallalophagy to phytophagy if the hypothesi
s is accepted that Camptodes is a member of the Oxycnemus group.