21.03.2015 Views

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

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

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

SHOW MORE
SHOW LESS

Create successful ePaper yourself

Turn your PDF publications into a flip-book with our unique Google optimized e-Paper software.

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

Hooray! Your file is uploaded and ready to be published.

Saved successfully!

Ooh no, something went wrong!