Cockroache; Ecology, behavior & history - W.J. Bell

estimate by Basset (2001) that cockroaches constitute approximately
24% of the arthropod biomass in tropical
tree canopies worldwide, and hints from various studies
suggesting that cockroaches may ecologically replace termites
in some habitats (Chapter 10). We address previously
unexplored aspects of their biology, such as the relationship
with microbes that lies at the heart of their
image as anathema to civilized households (Chapter 5).
As our writing progressed, some chapters followed unpredicted
paths, particularly evident in the one on mating
strategies (Chapter 6). We became fascinated with
drawings of male and female genitalia that are buried in
the taxonomic literature and that suggest ongoing, internally
waged battles to determine paternity of offspring. It
is the accessibility of this kind of information that can
have the most impact on students searching for a dissertation
topic, and we cover it in detail at the expense of addressing
more familiar aspects of cockroach mating biology.
We planned the book so that each chapter can be
mined for new ideas, new perspectives, and new directions
for future work.
An interesting development since Roth and Willis
(1960) was published is that the definition of a cockroach
Fig. P.1 A phylogeny of cockroaches based on cladistic analysis of 175 morphological and life
history characters; after Klass and Meier (2006), courtesy of Klaus Klass. Assignation of genera
to subfamilies is after Roth (2003c) and differs somewhat from that of K & M, who place Archiblatta
in the Blattinae and Phoetalia in the Epilamprinae. Pseudophyllodromiinae used here is
Plecopterinae in K & M. Based on their results, K & M suggest that Lamproblattinae and Tryonicinae
be elevated to family-level status. Mukha et al. (2002, Fig. 2) summarize additional hypotheses
of higher-level relationships. Phylogenetic trees of Vrs˘anský et al. (2002, Fig. 364) and
Grimaldi and Engel (2005, Fig. 7.60) include fossil groups. Lo et al. (2000), Klass (2001, 2003),
and Roth (2003c) discuss major issues.
is somewhat less straightforward than it used to be. Cockroaches
are popularly considered one of the oldest terrestrial
arthropod groups, because insects with a body plan
closely resembling that of extant Blattaria dominated the
fossil record of the Carboniferous, “The Age of Cockroaches.”
The lineage that produced extant cockroaches,
however, radiated sometime during the early to mid-
Mesozoic (e.g., Labandeira, 1994; Vršanský, 1997; Grimaldi
and Engel, 2005). Although the Carboniferous fossils
probably include the group that gave rise to modern
Blattaria, they also include basal forms of other taxa.
Technically, then, they cannot be considered cockroaches,
and the Paleozoic group has been dubbed “roachoids”
(Grimaldi and Engel, 2005), among other things. Recent
studies of extant species are also blurring our interpretation
of what may be considered a cockroach. Best evidence
currently supports the view that termites are nested
within the cockroaches as a subgroup closely related to
the cockroach genus Cryptocercus. We devote Chapter 9
to developing the argument that termites evolved as eusocial,
juvenilized cockroaches.
Roth (2003c) recognized six families that place most
cockroach species: Polyphagidae, Cryptocercidae, Noctixii
PREFACE