Cockroache; Ecology, behavior & history - W.J. Bell
Cockroache; Ecology, behavior & history - W.J. Bell
Cockroache; Ecology, behavior & history - W.J. Bell
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include Blaberus spp., which readily bury themselves<br />
in dirt or loose guano (Blatchley, 1920; Crawford and<br />
Cloudsley-Thompson, 1971), and Pycnoscelus spp., found<br />
in a wide variety of habitats as long as they can locate appropriate<br />
substrate for burrowing (Roth, 1998b; Boyer<br />
and Rivault, 2003). All stages of Pyc. surinamensis tunnel<br />
in loose soil, and are also reported from rodent burrows<br />
(Atkinson et al., 1991). The sand-swimming desert cockroaches<br />
fall into this category, as well as species such as Ergaula<br />
capensis, where females and nymphs burrow into<br />
well-rotted coconut stumps (Princis and Kevan, 1955), as<br />
well as the dry dust at the bottom of tree cavities (Grandcolas,<br />
1997b). Blattella asahinai is known to burrow into<br />
leaf litter and loose soil; they are sometimes pulled up<br />
along with turnips in home gardens (Koehler and Patterson,<br />
1987). Individuals of Heterogamodes sp. are known<br />
to bury themselves in sand or earth (Kevan, 1962). Several<br />
Australian species (Calolampra spp., Molytria vegranda)<br />
seem to spend the daylight hours underground,<br />
emerging to feed after dark (Rentz, 1996; D. Rentz, in<br />
Roth, 1999b). When collected during their active period<br />
or in light traps they usually sport sand grains on their<br />
bodies. In caves, Eu. posticus nymphs burrow in the surface<br />
of loose guano. They may be completely concealed,<br />
or may rest with their heads on the surface with their antennae<br />
extended up into the air. If the guano is compacted,<br />
the cockroaches remain on its surface and are attracted<br />
to irregularities such as the edge of a wall, a rock,<br />
or a footprint (Darlington, 1970). The recently described<br />
species Simandoa conserfariam congregates in groups of<br />
20 to 50 individuals of all ages deep within the guano of<br />
fruit bats; none have been observed on the surface (Roth<br />
and Naskrecki, 2003).<br />
Crevice Fauna<br />
The cockroaches considered crevice fauna are those that<br />
insert themselves into preexisting small voids in generally<br />
unyielding substrates. These include species found under<br />
bark, in bark fissures, in the bases of palm fronds and<br />
grass tussocks, in hanging dead leaves, empty cocoons,<br />
and hollow twigs, under logs and rocks, in piles of stones,<br />
rock crevices, and the excavated galleries of other insects.<br />
An example of the latter is the Malaysian cockroach Margattea<br />
kovaci, which lives in bamboo internodes accessed<br />
via holes excavated by boring Coleoptera and Lepidoptera<br />
(D. Kovach, pers. comm. to LMR). Burrowing<br />
and crevice-dwelling cockroaches can be categorically<br />
difficult to separate, particularly species that shelter under<br />
rotting logs, in rolled leaves, or in the litter wedged<br />
into the base of bunch grasses, spinifex, or the leaf axils of<br />
many plants. The spaces under rocks and stones are a particularly<br />
important microhabitat for cockroaches in unforested<br />
areas. Species of the genera Deropeltis and Pseudoderopeltis,<br />
for example, are abundant under the boulders<br />
“bestrewing the Masai steppe country” (Shelford,<br />
1910b). Rock-soil interfaces may also act as corridors between<br />
habitats, serving as oases for cockroaches moving<br />
between caves, or between patches of forest (Lawrence,<br />
1953). Some cockroach species are morphologically specialized<br />
to inhabit the wafer-thin crevices under bark or<br />
rocks (Fig. 1.10). The incredibly flattened bodies of tropical<br />
Australian Mediastinia spp. allow them to slip into the<br />
unfolding leaves of gingers, lilies, and similar plants during<br />
the day. At night they move to new quarters as the<br />
leaves of their previous shelters unfold (D. Rentz, pers.<br />
comm. to CAN).<br />
In Solid Substrate<br />
<strong>Cockroache</strong>s that excavate permanent burrows in solid<br />
materials such as wood or compacted soil are more specialized<br />
than those that use loose substrate or crevices.<br />
They typically exhibit a suite of ecological and <strong>behavior</strong>al<br />
features associated with their fossorial existence, and external<br />
morphology tends to converge. There are two major<br />
groups that fall into this category, the Cryptocercidae<br />
and the Panesthiinae, the latter of which includes the soilburrowing<br />
cockroaches. There are other species whose<br />
morphology suggests they are strong burrowers, but little<br />
has been published on their field biology. The hissing<br />
cockroaches, including Gromphadorhina portentosa, have<br />
the general demeanor of burrowers. In a recently published<br />
book on the natural <strong>history</strong> of Madagascar, however,<br />
the only mention of these cockroaches is as prey for<br />
some vertebrates and as hosts for mites (Goodman and<br />
Benstead, 2003).<br />
Burrows in solid substrates offer mechanical protection,<br />
as well as shelter from some classes of parasites and<br />
predators. The fact that dispersal in both the Cryptocercidae<br />
and Geoscapheini occurs following rainfall when<br />
excavation is likely to be more efficient (Rugg and Rose,<br />
1991; Nalepa, 2005) suggests that burrow creation is energetically<br />
costly. Pathogens may accumulate in tunnels,<br />
and occupants may not be able to escape if a predator enters<br />
the excavated space. It is unknown if burrowing cockroaches<br />
have strategies for dealing with flooded burrows,<br />
or with the often peculiar O 2<br />
to CO 2<br />
ratios that may occur.<br />
In Wood<br />
Dead wood is a tremendously diverse resource that varies<br />
with plant taxon, size (branch to bole), location (forest<br />
floor to suspended in canopy), degree and type of rot, orientation<br />
(standing versus prone), presence of other invertebrates,<br />
and other factors. Cockroach species from<br />
46 COCKROACHES