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DUANE D. MCKENNA, BRIAN D. FARRELL, MICHAEL S. CATERINO, CHARLES W. FARNUM,
DAVID C. HAWKS, DAVID R. MADDISON, AINSLEY E. SEAGO, ANDREW E. Z. SHORT,
ALFRED F. NEWTON and MARGARET K. THAYER.
Phylogeny and evolution of Staphyliniformia and Scarabaeiformia: forest litter as a stepping stone for diversification of nonphytophagous beetles.
Systematic Entomology, 2014.
Article first published online: 6 AUG 2014. DOI: 10.1111/syen.12093
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The beetle series Staphyliniformia exhibits extraordinary taxonomic, ecological
and morphological diversity. To gain further insight into staphyliniform
relationships and evolution, we reconstructed the phylogeny of Staphyliniformia
using DNA sequences from nuclear 28S rDNA and the nuclear protein-coding gene
CAD for 282 species representing all living families and most subfamilies,
a representative sample of Scarabaeiformia serving as a near outgroup, and
three additional beetles as more distant outgroups.
Under both Bayesian inference (BI) and maximum likelihood inference (MLI),
the major taxa within Staphyliniformia are each monophyletic:
(i) Staphylinoidea, (ii) Hydrophiloidea s.l., and the contained superfamilies
(iii) Hydrophiloidea s.s. and (iv) Histeroidea, although Staphylinoidea and
Hydrophiloidea s.l. are not strongly supported by MLI bootstrap.
Scarabaeiformia is monophyletic under both methods of phylogenetic inference.
However, the relative relationships of Staphylinoidea, Hydrophiloidea s.l. and
Scarabaeiformia differ between BI and MLI: under BI, Staphyliniformia and
Scarabaeiformia were sister groups; under MLI, Hydrophiloidea s.l. and
Scarabaeiformia were sister groups and these together were sister to
Staphylinoidea.
The internal relationships in Scarabaeiformia were similar under both methods
of phylogenetic inference, with Cetoniinae, Dynastinae?+?Rutelinae,
Hybosoridae, Passalidae, Scarabaeidae and Scarabaeinae recovered as monophyla.
Histeridae comprised two major clades: (1) Abraeinae, Trypanaeine and
Trypeticinae; and (2) Chlamydopsinae, Dendrophilinae, Haeteriinae, Histerinae,
Onthophilinae, Saprininae and Tribalinae. The relationships among
early-divergent Hydrophiloidea differed between BI and MLI, and overall were
unresolved or received only moderate to low nodal support.
The staphylinoid families Agyrtidae, Hydraenidae and Ptiliidae were recovered
as monophyletic; the latter two were sister taxa, and Staphylinidae?+?Silphidae
was also monophyletic. Silphidae was placed within Staphylinidae in close
relation to a subset of Tachyporinae. Pselaphinae and Scydmaeninae were both
recovered within Staphylinidae, in accordance with recent analyses of
morphological characters, although not always with recently proposed sister taxa.
None of the four major groups of Staphylinidae proposed by Lawrence and Newton
(1982) was recovered as monophyletic.
Certain highly specialized staphyliniform habits and morphologies, such as
abdominal defensive glands and reduced elytra, have arisen in parallel in
separate lineages. Further, our analyses support two major transitions to an
aquatic lifestyle within Staphyliniformia: once within Staphylinoidea
(Hydraenidae), and once within Hydrophiloidea s.l. (Hydrophiloidea s.s.).
On a smaller scale, the most common transition is from litter to subcortical
or to periaquatic microhabitats and the next most common is from litter to
carrion and to fungi. Overall, transitions to periaquatic microhabitats were
the most numerous. The broad picture in Staphyliniformia seems to be a high
level of evolutionary plasticity, with multiple possible pathways to and from
many microhabitat associations, and litter as a major source microhabitat for
diversification. In Scarabaeiformia, the most common transitions were from
litter to foliage, with flowers to litter, litter to flowers, and litter to
dung being next, and then litter to roots, logs or carrion. Litter is again
the largest overall source microhabitat. The most common transitions were to
foliage and flowers. It thus seems that the litter environment presents
ecological and evolutionary opportunities/challenges that facilitate entry of
Staphyliniformia and Scarabaeiformia into "new" and different ecological
adaptive zones.