i

AN OVERVIEW AND ANALYSIS OF GLOBAL ZOOGEOGRAPHY DISTRIBUTION

OF BLATTODEA

RYAN MITCHELL

Supervisor: Dr. Pamela Worrall BSc, PhD, PGCE.

A project submitted in partial fulfilment of the requirements for the Bachelor of

Science Degree in Animal Conservation and Biodiversity for the University of

Greenwich.

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DECLERATION

ACADEMIC SESSION: 2014-2015

PROJECT DECLARATION

“This project report is the result of the independent work of [name in full]. All

other work reported in the text has been attributed to the original authors and

is fully referenced in the text, and listed in the Reference Section”.

Student Name:-

Student Signature:-

Date:-

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AKNOWLEDGEMENT

I would like to thank and show my appreciation especially to Dr George Beccaloni for

letting me take on this project which I have thoroughly enjoyed. Without him and his

invaluable insight this project would have not been possible. From this project I have

come to appreciate cockroaches for what they truly are.

I would like to thank Dr Ben Price for proof reading my dissertation at a very stressful

period and the invaluable feedback which he provided. I would also like to thank him

for all the opportunities he has given me at the NHM.

I would also like to thank Dr Pam Worrall for becoming my Dissertation supervisor,

for providing advice, providing positive and constructive feedback, and making

herself available as much as possible for her students.

I also would like to thank my friends at Hadlow who supported me in the HE area

and kept me in a good mood throughout especially Paris Sadgrove, Alice Treharne,

David Courtneidge, Max Burford, and Michael James Lee.

Finally I would also like to thank my family for supporting me throughout the time of

writing my dissertation.

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ABSTRACT

Cockroaches are generally considered soil fauna their major role in the ecosystem is

the breakdown of complex carbohydrates (Bell et al, 2007), they also are vectors for

many species of fungi and bacteria (Fotedar et al, 1991). The majority of

cockroaches are known to be found in the tropics in study cockroaches biomass was

calculated at 24.3% higher than any other order (Basset, 2001). However they are

found in a wide variety of habitat across the world.

In this study the world distribution of cockroaches across the zoogeographic regions

was evaluated and analysed, the secondary data source was from the Cockroach

Species File (Beccaloni, 2013). This was used to see if there was a significant

difference between endemic species, species richness and genera richness between

zoogeographic regions (Holt et al, 2012). Also this study contains a world map of

species richness and is discussed in detail.

There was a significant difference between endemic species between zoogeographic

regions (T-test P=0.005), also provides evidence that the zoogeographic regions for

terrestrial invertebrates concept as endemic species percentages are very high in

the majority of the regions. The Nearctic region (52%) endemic species were as the

other regions have (74 %≥). This suggests the zoogeographic region boundary may

be not be positioned correctly in the case of cockroaches as the transitional zone is

very high and results in low endemism.

This study could be used to aid conservation and taxonomical efforts in specific

regions and countries which are highlighted in the study, with seemly unusually low

species richness and in some cases no data at all.

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Table of Contents

DECLERATION ....................................................................................................... ii

AKNOWLEDGEMENT ............................................................................................ iii

ABSTRACT ............................................................................................................. iv

Table of Figures ..................................................................................................... vii

1.0 INTRODUCTION .............................................................................................. 1

2.0 LITRATURE REVIEW ....................................................................................... 2

2.1 Morphology ....................................................................................................... 2

2.1 Colouration ........................................................................................................ 2

2.2 Sexual Dimorphism ........................................................................................... 3

2.3 Life Cycle .......................................................................................................... 4

2.4.0 Behaviour ....................................................................................................... 5

2.4.1 Solitary Behaviour .......................................................................................... 5

2.4.2 Social Behaviour ............................................................................................ 5

2.4.3 Parental care .................................................................................................. 5

2.4.4 Defensive behaviour ...................................................................................... 6

2.5. Taxonomy ........................................................................................................ 7

2.6 Evolution ........................................................................................................... 9

2.6.1 Termite evolution ........................................................................................... 9

2.7 Ecology ........................................................................................................... 10

2.7.1 Forest ecology ............................................................................................. 11

2.7.2 Desert ecology ............................................................................................. 12

2.7.3 Aquatic Habitats ........................................................................................... 14

2.7.4 Predators ..................................................................................................... 14

2.7.5 Contribution to ecosystem services ............................................................. 15

2.7.5.1 Nitrogen fixation ........................................................................................ 15

2.7.5.2 Methane cycle ........................................................................................... 15

2.7.5.3 Pollination ................................................................................................. 15

2.7.5.4 Waste removal .......................................................................................... 16

2.8 World species Distribution ............................................................................... 16

2.9 Wallace and zoogeographical regions ............................................................ 17

2.10 Endemism and species richness ................................................................... 19

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2.11 Cockroach and Conservation ........................................................................ 19

2.12 Contributors to cockroach taxonomy and research ....................................... 20

2.12.1 Robert Walter Campbell Shelford .............................................................. 20

2.12.2 Karl Richard Hanitsch ................................................................................ 20

2.12.3 Dr Louis M. Roth ........................................................................................ 20

2.14 Lack of research Insect data ......................................................................... 21

3.0 AIMS AND OBJECTIVES ............................................................................... 21

3.1 Hypotheses ..................................................................................................... 22

3.1.1 Relationship between species richness and zoogeographic region ............. 22

3.1.2 Relationship between Genus richness and Zoogeographic region .............. 22

3.1.3 Relationship between endemic species and Zoogeographic Region ........... 22

3.1.4 Relationship between Genera richness and Species richness in relation to

zoogeographic region ........................................................................................... 22

4.0 METHODOLOGY ............................................................................................ 23

4.1.1 Data collection ............................................................................................. 23

4.1.2 Data analysis ............................................................................................... 24

4.2 Data statistical analysis ................................................................................... 25

4.2.1 Relationship between endemic species and Zoogeographic Region ........... 25

4.2.2 Relationship between Genus richness and Zoogeographic region .............. 25

4.2.3 Relationship between Species richness and Zoogeographic region ............ 25

4.2.4 Relationship between Genera richness and Species richness in relation to

zoogeographic region ........................................................................................... 25

4.3 RESULTS AND STATISTICS ......................................................................... 26

5.0 Discussion ....................................................................................................... 31

5.1 Percentage of Endemic species by zoogeographic region.............................. 31

5.2 World distribution map of species richness ..................................................... 32

5.3 Blattodea hotspots .......................................................................................... 35

5.4 Species richness by Zoogeographic region .................................................... 35

5.5 Relationship between Genus richness and Zoogeographic region ................. 37

5.6 Relationship between Genera and Species richness by Zoogeographic region

.............................................................................................................................. 37

5.8 How the study could be improved ................................................................... 38

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6.0 CONCLUSION ................................................................................................ 38

7.0 FURTHER RESEARCH .................................................................................. 40

8.0 REFRENCE LIST ............................................................................................ 41

9.0. APPENDICIES .................................................................................................. 48

9.0.1. Appendix 1 .................................................................................................. 48

9.0.2 Appendix 2 ................................................................................................... 54

Table of Figures

Figure 1 Sexual Dimorphism (Roth, 1991) ................................................................. 3

Figure 2 Life stages of Methana marginalis (Roth, 1991) ........................................... 4

Figure 3 Taxon Tree (Van der Wart, 2015) ................................................................ 7

Figure 4 Family taxonomy of the suborder Blattaria (Roth, 2003) .............................. 8

Figure 5 the addition termite sister group (Inward et al, 2007) ................................. 10

Figure 6 map of the world with tropical rainforest highlighted (Britannica, 1997) ..... 12

Figure 7 map of the world with the desert regions highlighted (Hendrix et al, 2014) 13

Figure 8 zoogeographic regions map and key (Beccaloni, 2014) ............................. 17

Figure 9 A revised world map of zoogeographic regions (Holt et al, 2013) .............. 18

Figure 10 Excel document master list ...................................................................... 23

Figure 11 Species richness by country ..................................................................... 24

Figure 12 World cockroach species richness by zoographical region ...................... 26

Figure 13 World cockroach genera richness by zoographical region ....................... 27

Figure 14 World Cockroach species and genera percentage comparison by

zoogeographic region ............................................................................................... 28

Figure 15 Species endemic percentage by zoogeographic region ........................... 28

Figure 16 Biodiversity hotspots of Blattodea of species diversity ............................. 29

Figure 17 World map of cockroach species richness by political region .................. 30

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1.0 INTRODUCTION

Cockroaches are a relatively small order of insect with 4549 species (Cockroach

species file, 2014); they are generally considered pests and have a poor public

image. Cockroaches are an understudied order in the entomological field; they are a

highly diverse group they are found across the world in a large variety of different

habitats. The tropical rainforests are regions know for the highest cockroach diversity

(Bell et al, 2007).

Their main role in the ecosystem is they are responsible for breaking down

complex carbohydrates and extract minerals in decaying material back into the

ecosystems (Lavelle, 2002). They are also renowned for their ability to transport

bacteria, fungi and as vector, transporting these taxon species in leaf litter could

enhance and increase the nutrient recycling process (Fotedar et al, 1991).

There is a lack of knowledge about the global distribution and zoogeography of

insects in general with an estimate only 30% of arthropods are known (Scheffers,

2012). Only 25 species of cockroach are acknowledged on the IUCN red list and 15

of them are considered endangered or critically endangered (ICUN red list, 2014).

The conservation of cockroaches relies on knowledge of species populations and

species richness within a habitat or region.

The aim of the study will be able to assess the currently known species and genus

richness of zoogeographic regions. To evaluate species endemism in each

zoogeographic region to locate possible region which maybe phylogenetic distinct.

To produce a map of the world species richness by country and to critically

discussed to highlight areas of particular interest to enhance the study. The

secondary data in which the study is based is from the online source the cockroach

species file.

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2.0 LITRATURE REVIEW

2.1 Morphology

Cockroaches are classified under the Class of Insecta they are readily identifiable

from there features 3 pairs of legs, three body parts head, thorax and abdomen and

one pair of antenna. The Class contains 25-30 orders which are found worldwide

(bugguide, 2015). The cockroach distinct features such as 10 segments of tergum

which are located upon the abdomen, other features such as two pairs of wings

(these are not present in all species) with the front pair are mesothoracic and are

called tegmina they dark and leathery in appearance have a protective. The hind

wings are membranous and transparent; there function is for flight these wings are

protected by the front tegimina. The Head is comprised of compound eyes which are

usually fully developed, however they are reduced or absent in cave dwelling

species. The antenna are many segmented and filament like, the mandibles are

strong and contain teeth. The thorax has a prothorax with a shield like pronotum

which occasionally overlaps the head (Roth, 1991). The pronotum it has become

adapted in many species into various tools such as a wedge, plug and ram, the

majority of species where the pronotum overlaps the head belong to digging species

(Bell et al, 2007).

The typical flat body shape of the cockroach is due to its ecological niche, this is

functionally related to their microhabitats in which they exist. Their body shape

allows them to occupy narrow and horizontal extents found within leaf litter and tree

crevices. Pancake syndrome is the typical look for many species this is where the

species is dorsoventrally compressed this gives protection against both abiotic and

biotic factors such as predation against ants (Mackerras, 1967b).The cockroaches

can become immobile by clamping down to a trees surface protecting their

vulnerable undersides and water retention in dry conditions (Grancolas et al, 1994).

2.1 Colouration

Cockroach colouration is determined by behavioural ecology such as Crypsis or

aposematic behaviour see (2.4.5 Defensive behaviour), the majority of species

possess dull and dark colours which are appropriate for their cryptic lifestyle within

decaying plant material and nocturnal behavioural activities. However it has been

found that a few species colouration changes due to environmental factors within the

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ecosystem, the species Ectobius panzeri found in Britain are considerably darker in

higher altitude populations and females often turn darker in colour during the end of

breeding season (Brown, 1952). The species Panchlora nivea colouration changes

in their life stages as their behaviour changes the nymphs are browns where as in

the adult stage they are bright green (Roth and Willis, 1958b). The diurnal species

are generally more conspicuous they have three broad categories small colourful

species, aposematic species and Batesian mimics (Bell et al, 2007).

2.2 Sexual Dimorphism

Sexual dimorphism is present in many cockroach species, there are some species

this occurs as size, colouration or wing presence this can become extreme and

males and females appear to be different species see (Figure 1). There is a variety

of sexual dimorphic characteristic features in cockroaches. The different

morphological features in sexual dimorphism suggests that the demands of winning

a mate are highly determined upon they influence the morphological evolution of the

species (Roth, 1991).

Figure 1 Sexual Dimorphism (Roth, 1991)

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2.3 Life Cycle

Figure 2 Life stages of Methana marginalis (Roth, 1991)

A cockroach’s lifecycle is comprised of three stages see Figure 3. Egg (ootheca) (A),

nymph (B, C, D) and Adult (E) this is known as incomplete metamorphosis or

hemimetabolism (Gullan, 1994). The eggs are deposited in groups they are enclosed

inside an ootheca which is a produced by the female. The ootheca is either carried

externally in oviparous species or internally in viviparous species. The oviparous

species tend to produce oothecas which are complete and generally quite resistant

to desiccation. However female viviparous species produce usually more flexible

walled ootheca’s with absent or simple keels (Cochran, 1999).

Once the cockroaches emerge from the ootheca, this is known as the nymph stage

of their lifecycle. They generally resemble the adults of their species although lack

wing structures and usually differ in colouration. They moult as they grow the

number of moult varies between taxa and sometimes differs between sex within the

same species (Roth, 1991). The moulting process is required as their rigid cuticle

restricts growth at each stage. The stages between moults are known as instars. The

outer cuticle periodically sheds as the cockroach in order for the cockroach to grow

and reach the imaginal instar or adult stage (Gullan, 1994).The most noticeable

change in the imaginal instar is usually the appearance of wings taxa and sex

dependent.

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2.4.0 Behaviour

2.4.1 Solitary Behaviour

There is low number of truly solitary species of cockroaches that only meet other

members of the species in order to mate. Deep cave cockroaches come into this

category however they do occasionally congregate around food sources but they

have large ranges and only meet another cockroach in order to reproduce

(Langecker, 2000). Thanatophyllum akinetum is a species which exists in French

Guyana that has evolved a cryptic solitary lifestyle. This has allowed the species to

co-exist with army ants as they actively keep a set distance from other individuals

and spend their time motionless in the leaf litter avoiding attention from their

predators (Grandcolas, 1993).

2.4.2 Social Behaviour

The genera Gromphadorhina Madagascan hissing cockroaches live in social

systems they must be able to recognise dominate group members to avoid costly

contests between members. The males most noticeably have been observed to

show display behaviours such as thorax thrashing to reduce the chance of fighting,

they use also complex acoustic behaviours hence the common name. The social

system is based on dominance based territorial behaviour for mate choice, the males

are sexually dimorphic they have pronotal horns which are used in fighting

behaviour. The males appear to show associations with rank of individuals in the

hierarchy (Clark, 1993).

2.4.3 Parental care

Majority of cockroaches show a form of parental care this is a behaviour that

promotes survival, growth and development of nymph cockroaches. Cockroaches

are considered one of the largest groups of insects that show parental care

behaviours (Hinton, 1981). In both oviparous and viviparous female cockroaches the

embryos are protected and provisioned within the body of the cockroach. In

oviparous species the adults exhibited behaviour such as production of oothecal

case, the preparation of ootheca deposition site, concealment of ootheca and the

defending behaviour of protecting the ootheca. Brooding behaviour has also been

6

observed in oviparous females, this is where the nymphs cluster on and around the

female cockroach for periods of time after emergence (Grancolas, 1993).This

behaviour was witnessed for less than a day although this was observed in

laboratory conditions. This brooding behaviour is believed to protect the nymphs as

they are most vulnerable after hatching as their cuticle is soft for several hours and in

this state they are at high risk from predators and cannibalism. Another factor could

be the transferring of gut microbiota from the adult female to the nymphs through a

faecal meal, however there is currently a lack of research in this area to support the

brooding behaviour (bell et al, 2013).

Sub social behaviour on extreme form of this is shown by Byrostria fumigata the first

instars are able to recognise their mother and show a preference of aggregating

beneath the mother cockroach for the first 15 days after hatching. Other forms of

sub social behaviour these include species with adaptations in their morphology of

nymphs or the females facilitate care. These adaptations include appendages on

nymphs to help cling on to the female. Adaptations to mouth parts to facilitate unique

feeding behaviours such as the species Thorax porcellana, the nymphs obtain

nutrients from the mother with their specialised sharp mandibles. They pierce the

cuticle on her abdomen to feed on her haemolymph (Beccaloni, 2013). Some

species have evolved external brooding chambers under their wing cases (Bell et al,

2007). Species of the Perisphaerus genus have developed the ability to

(conglobulation) roll into a ball like a pill millipede to protect ventrally clinging

nymphs. The nymphs are born blind and have tubular mouth parts which are unique

in cockroaches; these mouth parts fit into appendages within the mothers mid and

hind legs. The nymphs are believed to extract nutrients from these pores; the

nymphs do not develop functioning eyes until the third instar (Beccaloni, 2013).

2.4.4 Defensive behaviour

Cockroaches are prey for a variety of arthropods species and numerous

entomophagous vertebrates, amphibians, birds and mammals. Cockroaches have

developed species specific defences such as aposematic mimicry of Coleoptera

especially coccinellids and chrysomelids however the majority of models are not

known (Wickler, 1968). Another example of aposematic mimicry is the species

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Cardacopis shelfordi have the yellow and black colouration which mimics dangerous

hymenoptera species (Roth, 1988).

Other defence mechanisms are chemical secretions many of the polyzosteriinae

family share this method of defence they produce aliphatic compounds. Some

species of this family have developed a warning stance which they angle their bodies

vertically revealing bright warning colouration on their coxae and venters. The gland

opening is faced toward the threat if the warning stance is not effective it will

discharge its chemical defence, 2-methylbutanal is a chemical commonly expelled

(Roth, 1991).

2.5. Taxonomy

Figure 3 Taxon Tree (Van der Wart, 2015)

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Cockroaches are in the higher classification the order Orthoptera. Cockroaches are

primitive orthopteran insects and are more generally placed into the suborder

Dictyoptera with Mantodea and Isoptera (Roth, 2003). Mckitterick developed a

taxonomic system which 4 morphological systems; female genetalia, male genetalia,

proventriculus and oviposition behaviour. From the studies conducted the conclusion

was there are 2 subfamilies evolved along two divergent lineages and have been

grouped into two superfamilies Blattoidea; comprised of two families Cryptocercidae

and Blattidae (Mckitterick, 1964). The other Blaberidae consist of the families

Polyphagidae, Blattellidae and Blaberidae, 20 subfamilies extend from these

superfamily lineages see (Figure 4) (Roth, 2003).

Figure 4 Family taxonomy of the suborder Blattaria (Roth, 2003)

Taxonomical issues with Blattodea especially in the superfamily Blattaria in relation

to the homogeneity of their morphological features. Some author’s species

descriptions which were used as diagnostic evidence were unnecessarily brief, when

describing morphological features or characters. This has now been discovered that

these features have no or little significance of species level identification, It has also

been noted that some species descriptions were so ambiguous it could relate to

other species and in some instances other genera (Fallia et al, 1987).

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2.6 Evolution

Cockroaches are one of the earliest known group of winged insects; the first fossil

records date back to the carboniferous period 359.2 to 299 million years ago

(Kamhampati, 2005). There are numerous journals the Blattodea on prehistoric

species collected from fossil resins; these are mainly nymphs rather than much rare

adult specimens. However the taxonomic information from resin nymphs is no less

important than provided from adult impressions in rocks which contain wing venation

information (Gorokhov, 2006).

2.6.1 Termite evolution

Termites are believed to have evolved from the genus Cryptocercus which are a

group of wood feeding cockroaches as both shares symbiotic gut flagellates with

early evolved species of termites. The nymphs of both groups share similar

morphological features, the evolution occurred when nymphs of the Cryptocercus

required longer parental care as they are dependent on the transfer of gut flagellate

transfer through proctodeal trophallaxis. This evolutional adaptation to a eusocial

behaviour in termites eventually led to the loss of the ootheca, this was due to the

nesting behaviour this provided both protection and its regulated climate which made

it unnecessary to waste energy producing a ootheca. See (Figure 5) this is the

suggested placement of termitidae sister group into Blattodea order (Inward et al,

2007).

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Figure 5 the addition termite sister group (Inward et al, 2007)

2.7 Ecology

Cockroaches have developed their stereotypical flattened shape due to their crevice

inhabiting niche within an ecosystem. This allows them to exploit narrow, horizontal

and spaces found within in ecosystems such as decaying leaf litter. Even distantly

related species occupy a very similar ecological niche however they differ through

changes of similar morphological features which reflects the demands of their

surrounding environment (Bell et al, 2007).

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The majority of cockroaches are considered soil fauna because of their diet is

dependent on decaying matter of animals and plants. They occupy a range of niches

within the soil ecosystem e.g. upper litter horizon, burrowing species can be found in

the mineral soil level, other species live within the suspended soil (Eienbeis et al,

1985). Cockroaches on a global scale are responsible for breaking down complex

carbohydrates and mineralizing nutrients of plant decaying matter in all ecosystems

(Lavelle, 2002).

The cockroach has an important role as vectors of bacteria and fungi they are moved

from to different locations via the cuticle (Fotedar et al, 1991). However cockroaches

generally have clean external surfaces they have been observed grooming and

cleaning there extremities with their modified comb like legs up to 50% of the time.

This grooming behaviour is believed to be related to prevent disease spread within

their habitat; the bacterium which is digested might also have another benefit in

species which have gut bacteria which neutralize ingested pathogens (Arnold, 1974).

In species that allogroom in their development stage there is evidence to suggest

there is a nutritional reward for the groomer, this behaviour in social species also

keeps other individuals within the group free of infections (Dhanarajan, 1978).

2.7.1 Forest ecology

In a temperate forest environment cockroaches are usually restricted to a minor

niche within the soil biology due to low population densities such as ectobius sp in

European woodlands (Bell et al, 2007). However in samples collected from four

forest types in Gunug Mulu in Sarawak, cockroach biomass was recorded at 43% of

the invertebrate biomass in alluvial forest, 33% in Dipterocarp forest, 40% in Heath

forest and 2% in a forest located on limestone. The tropical rainforest however

provides many niches within the canopies and arboreal habitats, in a study that

reviewed arthropod composition of the canopy 5.3% of individuals collected where

cockroaches however when represented in biomass composition they were found to

represent 24.3% with hymenoptera second which was comprised mainly of

formicidae (Basset, 2001). The Rainforest canopy is a relatively difficult environment

to survive with high exposure to sunlight and radiation, also other factors such as

wind and heavy rainfall are common occurrences (Madigosky, 2004). The Central

American species Epilampra irmleri their ecological decomposition rate has been

12

quantified to consume an estimated 6% of the annual leaf litter in the inundation

forests (Irmler et al, 1979). See (Figure 6) show tropical rainforest regions around the

world this is where cockroach species richness is predicted to be highest (Bell et al,

2007).

Figure 6 map of the world with tropical rainforest highlighted (Britannica, 1997)

2.7.2 Desert ecology

Deserts excluding the Polar Regions cover around 25% of the earth’s surface,

deserts are defined by the amount of precipitation they receive on an annual basis.

By definition a desert is an area of region which receives less than 25cm of rainfall a

year (Hendrix et al, 2014). Figure 7 below shows a map of the world with the desert

regions highlighted and labelled. There are a number of cockroach species which

have adapted to more arid environments such as deserts. Deserts generally have

low and unpredictable precipitation, extreme contrasting temperatures and low soil

organic matter (Wall et al, 1999). The family Polyphagidae has the most desert

dwelling species compared to the other families, there are some species of blattellids

and a small number of blattids (Bell et al, 2007).

13

The behaviour of cockroaches in desert environments have evolved to survive by

‘sand swimming’, which is the ability to burrow quickly below the sand surface.

Moisture in deserts is found beneath the sand at a depth of 30-60cm also

temperature declines as with depth. This area is where cockroaches are

predominantly found in the day as the microclimate is favourable, with reduced

temperature and higher humidity levels prevents excess water loss and overheating

(Kinlow, 1999). During the night cockroaches’ move towards the surface to feed on

decaying leaves, roots and various other food sources, however they still remain

under the sand to avoid predators such as scorpions (Bell et al, 2007). The Sand

roach the genus Arenivaga this group of Polyphagidae has the ability to

osmoregulate to absorb water from the surrounding environment this is unusual due

to the size of the cockroach 800mg which is relatively large whereas most

osmoregulatory species are much smaller (Edeny, 1974). Around half of the known

deserts species are found within microhabitats created by vertebrate species such

as ground squirrels, gerbils and tortoises. However some species are not restricted

to these habitats as they are found across a variety of micro habitats e.g. Arenivaga

floridensis has been observed in mice burrows, loose sand and in vegetation

communities of scrub (Atkinson et al, 1991).

Cockroaches in deserts are known to contribute the breakdown of organic matter

and have an impact on the nutrient flow within this ecosystem. Anisogamia

Figure 7 map of the world with the desert regions highlighted (Hendrix et al, 2014)

14

tamerlana is a species found in Turkmenistan they consume around 17% their dry

body mass daily. They are known to improve soil fertility the faecal matter they

produce has up to 10 times the nitrogen content then their food source (El-Ayouty et

al, 1978). In habitats where other decomposers are less abundant such as millipedes

and earthworms cockroaches have filled the niche such as the Australian species

Dezmososteria cincta in the breakdown of eucalyptus leaf litter (Rentz, 1996)

2.7.3 Aquatic Habitats

There are two categories of aquatic cockroach species phytotelmata aquatic habitats

which exist on or within plants such as bromeliads, and species which exist in ponds,

rivers and streams. Cockroaches live at the surface of the water however submerge

to escape predators or to hunt for food. Sixty-two species alone have been found

within bromeliads however it is unknown how many are restricted and are dependent

on the microhabitat further research should be carried out in this area to help aid any

future conservation efforts (Roth et al, 1960). Many species of the Epilamprinae live

along streams and pool habitats they feed upon decaying vegetation around the

water’s edge.

Many aquatic cockroaches have adapted their behaviour to submerge into the water

when disturbed by movement or shadows this allows them to escape from possible

predators. They cling on to the vegetation or rocks under the water’s surface for up

to 15 minutes (Bell et al, 2007). In many aquatic species the abdomen tip has been

adapted to a snorkel; also trapping an air bubble has been observed to maintain an

air supply whilst submerged (Wiedner, 1969).

2.7.4 Predators

Cockroaches as a prey item feed a vast array of species such as pitcher plants

Nepenthes sp (Roth and Willis, 1960). There is evidence to show that the genus

Parcoblatta in South Carolina is the one of the main prey items of the endangered

Picoides borealis Red cockaded woodpecker (Horn and Hanula, 2002). Another

15

endangered species which is dependent on cockroaches as a food source is the

Copsychus sechellarum the magpie robin on the Seychelles (Komdeur, 1996).

2.7.5 Contribution to ecosystem services

2.7.5.1 Nitrogen fixation

Cockroaches potentially have influences into the biogeochemical cycles such as the

Nitrogen the Cryptocerus genus are the only known species of cockroach which

contain microbes which are able to fixated atmospheric nitrogen (Slaytor, 1992).

These species have a nitrogen fixation rate which equates to similar rates as

termites on a body weight basis. The ecological significance is currently unknown,

however where they occur in high prevalence such as the montane mesic forest the

Cryptocerus is the dominate wood feeding species (Bell et al, 2007).

2.7.5.2 Methane cycle

Methane seems to be a universal by-product because of the methanogenic bacteria

which is found within their hindgut (Hackstein, 1994). This occurs in tropical species

of cockroaches regardless of derivation, during laboratory testing in resting

cockroaches found methane, carbon dioxide and water where to be released

simultaneously suggesting the gases are respired. However due to variable such as

cockroach age and fibre content it is difficult to estimate methane production on a

global scale. It has been suggested that the cockroaches make a significant

contribution to atmospheric methane particularly in the tropic regions (Gijzen et al,

1992).

2.7.5.3 Pollination

Some cockroaches are found feeding upon nectar and pollen, in the forests of Japan

the adult Margattea sumana visit nectaries at the base of egg like inflorescences of

the root parasite balanophora sp. The visits coincided with the evening nectar

secretion, the pollen grains were observed to be attached to the tarsi and mouthparts

although the crosspollination success has yet to be measured assumptions of low

success rates in this species have been made because it’s a flightless species (Bell

et al, 2007). There is stronger evidence of cockroaches being effective pollinators in

the lowland mixed dipterocarp forest in Borneo, this area has low bee densities in the

16

canopies. Blattellid cockroaches where observed feeding on pollen of the species

Uvaria elmeri a member of the Annonaceae family. The pollen grains were found

both within the gut and on the surface of the head, and the timing of visits coincided

with nocturnal dehiscence of anthers. Therefore due to the lack of bees it has been

suggested that the cockroaches may have evolved to fill the ecological niche of the

pollinators (Basset, 2001).

2.7.5.4 Waste removal

Many cockroach species niche is feeding upon faecal and waste matter of a large

variety of various taxa. The species found in microhabitats such as bird nests,

mammal burrows and social insect nests provide sanitation services which opposes

their reputation of carrying diseases. Evidence suggests the Parcoblatta nymphs

prolong the colony cycle of Vespula squamosa, they remove debris, as well as

keeping bacterial and fungal population in low densities (Macdonald et al, 1983).

2.8 World species Distribution

World species distribution is not evenly spread across the world there are areas with

higher species richness and higher levels of endemism compared to other regions

(Mittermeier, 2008). Currently 44% of the world’s known species diversity is

endemic to 12% of the earth’s surface in 25 different locations which are known as

Hotspots (Kitching, 2000).

The total number of species and distribution data is essential to develop an

understanding of the species richness and patterns of biodiversity within

ecosystems, without this knowledge the future and conservation of these species

(Dolphin et al, 2001). To estimate world distribution maps can be difficult due to

completeness of global inventories for example estimates suggests mammals

around 97% known where as 30% in arthropods. Taxonomists are evenly

distributed through taxa however this is a disadvantage because there are over 100

times more invertebrates than vertebrates. Another underlying issue are the where

about of unidentified species this is a concern because of current habitat destruction

rates the chances of these species being discovered before their habitat is

17

destroyed is questionable (Scheffers, 2012). There has been no study in this field

for specifically cockroaches or invertebrate distribution in general on a world basis.

2.9 Wallace and zoogeographical regions

Alfred Russel Wallace was the joint publisher of the theory of evolution by natural

selection with Charles Darwin in 1858. He was a field biologist and specimen

collector who spent almost 8 years in the Malay Archipelago where he collected

110,000 insects, 7500 shells, 8050 bird skins and various other curiosities. During

his time there he wrote and published a book on his journey The Malay Archipelago

which became one of the most celebrated travel writings of the 19th century

(Beccaloni, 2008). In 1876 Wallace published a map of the global terrestrial

zoogeographical regions which was theorised upon both distribution and taxonomic

relationships between various taxon groups (Holt et al, 2012). Below shows a basic

map of the world divided into these 9 regions (Figure 8) is a key to colouration of the

zoogeographical regions.

Figure 8 zoogeographic regions map and key (Beccaloni, 2014)

Zoogeographic regions are hypothesised upon geological tectonic events which

occurred during the Mesozoic era where the continents Gondwana and Lauraisa.

Madagascar is a prime example of where geological isolation has driven evolutionary

distinctiveness. However new evidence suggests that environmental boundaries

which existed for long periods also act as historical barriers , Examples of

18

environmental barriers temperature and humidity Nearctic and Neotropical regions

(Proches, 2012).

Figure 9 A revised world map of zoogeographic regions (Holt et al, 2013)

A study was carried out on global phylogenetic and distribution relationships

between 21,037 vertebrate species this included amphibians, birds and mammals.

From this study it was concluded there where 11 larger zoogeographic regions the

new suggested regions are Sino-Japanese, Sharo-Arabian and Panamanian see

Figure 9 above (Holt et al, 2013).

This study was carried out on terrestrial vertebrates and not invertebrates this must

be taken into consideration for global zoogeographic studies as there maybe

difference in phylogeny and distribution relationships in comparison to vertebrate

groups. As many invertebrate species may have evolved earlier changing the

cladistics and how they have become distributed. The paper suggested the addition

of invertebrate data could affect the regions distribution (Holt et al, 2013) Only 9

regions will be included in this study as that is how the data is arranged and is

difficult to change into the new perceived system as no specific locality data is

included in the data sets. These 9 regions are used in various entomological

collections such as the as the Natural History Museum London as a method of

having zoogeographic distribution data easily displayed on draws as a colour code

system (Scoble, 2002).

19

2.10 Endemism and species richness

Endemic in an ecological term is a species which is unique to a geographic location,

nation or habitat and found nowhere else (Hogan, 2011). Endemic richness is a

fundamental to aid global conservation as understanding the levels of endemism can

prioritise areas which may previously be overlooked form a conservation protection.

Islands are known for high levels of endemism however there species richness is

known to be lower than the mainland, analysing both levels of endemism with

species richness can identify specific regions which are high in both and should be

priorities for conservation efforts (Keir, 2009). The traditional method of providing

evidence for conservation protection is species and genera richness the taxonomy is

assumed to be equivalent unit, Genera is preferred over species as generally there

are less taxonomic issues at this level (Perez-Losada, 2003). The loss of a species

with no or limited related extant species in comparison to a species with many

related species would be a much greater loss to genetic and phylogenetic diversity

and should be conservation efforts should prioritised over the other species.

2.11 Cockroach and Conservation

Cockroaches live in many areas where there are large conservation threats to

biodiversity such as deforestation, urbanisation, global warming and agricultural

practices. Cockroaches are not high on the agenda for invertebrate conservation as

they are not a popular or well-studied group of insects (Bell et al, 2007).There are 25

species of cockroach found on the IUCN redlist, all these species where assessed

by Gerlach and reviewed by Hochkrich (IUCN, 2015).

A quick breakdown of the 25 species they are all found in the Seychelles region.

One species is now considered extinct Margetteoidea amoena and has not been

seen in the wild since 1905. 8 species are critically endangered, 7 species

Endangered, 1 species data deficient and 8 least concern.

The Desroaches cockroach Delosia ornata a critically endangered species found

endemic to Desroaches Island. The population was last estimated in 2006 with less

than 300 adults and 600 nymphs and is threatened due to hotel development. The

species is currently surviving in woodland fragments <100m2, the island highest

20

point is less than 1m and is all so under threat of rising sea levels. There is currently

no conservation plan put in place for this species (Gerlach, 2012).The other species

Gerlach cockroach Nocticola gerlachi is located on the Seychelles granitic islands

which has a status of endangered it is located to a site of 10km2 of costal gardens

which is a secure habitat however at risk from rising sea levels (Gerlach, 2012).

2.12 Contributors to cockroach taxonomy and research

2.12.1 Robert Walter Campbell Shelford

Shelford was an entomologist who was born in Singapore in 1872; he was a curator

at the Sarawak museum for 7 years then became assistant curator at the hope

entomological collection in 1905. He published papers on a variety of orders

Lepidoptera, Orthoptera and Phasmantodea. He published his first paper on

cockroaches in 1906 (studies of the blattidae. Transactions of the entomological

society of London, 1906: 231-280) this paper described 38 species and 5 genera. In

8 years Shelford published 27 papers and described 284 new species, since only

13% of them have been synonymised this indicates his studies where thorough

(Cockroach Studies, 2006).

2.12.2 Karl Richard Hanitsch

Hanitsch was born in Germany in 1860; also studied other groups of organism like

Shelford did these included molluscs, sponges and Lepidoptera. He was a curator at

the raffles museum in Singapore and in 1919 became an associate curator at the

Hope entomological collections. In 1915 he produced his first Blattidae paper

(Malayan Blattidae) he only described 9 species, by the end of his career he

produced 36 papers on cockroaches and described 300 species. Since only 10% of

the species he described have been synonymised. Hanitsch contributed the most

specimens to the HEC collections compared to everyone else, with 162 type

specimens around 28% of the collections type material (Cockroach Studies, 2006).

2.12.3 Dr Louis M. Roth

Roth was a researcher at Harvard University he started his career studying

mosquitos for 30 years, whilst in Natick he became interested in cockroaches

21

because of their importance to the U.S military and the ease of rearing them. He

devoted his time into studying their behaviour and physiology; He was given an

office at Harvard University’s museum of zoology. Whilst at Harvard he worked on

cockroach taxonomy and described 400 species previously undescribed that he

collected from field trips and received in from researchers around the world. He also

named 20 genera and published over 40 papers during the 30 years he studied

cockroaches. He received an award for his achievements in insect systematics the

Thomas say award in 1995.He continued his research into his retirement and often

worked 7 days a week without pay his affection for cockroaches was truly admirable

(entsoc, 2012).

2.14 Lack of research Insect data

There are large gaps of knowledge in invertebrate global biodiversity this is generally

because there is a lack of comprehensive data sets of these orders these data sets

are generally available in vertebrates and plants in which most of world speciation

and zoogeography has been based upon and where research has predominantly

carried out (Economo et al, 2014).

3.0 AIMS AND OBJECTIVES

There has been no study in this area of cockroaches this is because the data until

recently has been difficult to obtain. The aim of the study will be able to assess

currently the known species and genus richness of zoogeographic regions, to assess

species endemic richness of zoogeographic region. To produce world map species

richness by political region this will indicate countries with high species richness and

ones with low species richness. From the data analysis of Blattodea world

distribution, assess areas for possible research and provide evidence to theories of

cockroach ecology and aid research for cockroach conservation.

22

3.1 Hypotheses

3.1.1 Relationship between species richness and zoogeographic

region

H1- There will be a significant difference between species diversity and

zoogeographic region

H0- There will be no significant difference between species diversity and

zoogeographic region

3.1.2 Relationship between Genus richness and Zoogeographic

region

H1- There will be a significant difference between genus distribution and

zoogeographic region

H0- There will be no significant difference between genus distribution and

zoogeographic region

3.1.3 Relationship between endemic species and Zoogeographic

Region

H1- There will be a significant difference between endemic species and non-endemic

species in each zoogeographic region

H0- There will be no significant difference between endemic species and non-

endemic species in each zoogeographic region

3.1.4 Relationship between Genera richness and Species richness

in relation to zoogeographic region

H1- There will be a significant difference between genera richness and species

richness in relation to zoogeographic region.

H0- There will be a no significant difference between genera richness and species

richness in relation to zoogeographic region

23

4.0 METHODOLOGY

For this study secondary data was used the Blattodea species world distribution

database on the Cockroach Species File online data in the form of an excel sheet

world valid cockroach species master file produced by Dr George Beccaloni.

4.1.1 Data collection

The secondary data is collected from various specimens from institutions around the

world such as the Natural History Museum London; this study’s data was collected

from the Cockroach species file which is a website and online database. This online

database has been created for international use for taxonomic study and online

management of the world’s cockroach data. This contains detailed information on

each species which has been data based into the system by various individuals who

are taxonomists and administrators of the website. This information has been

manually inserted into a Microsoft excel document which can be used to create an

appropriate table for the data this includes (Distribution, species, genus, Family,

original taxonomic name, author, year described) see (Figure 10) below once all the

data has been collected and input into excel document as a master list then then the

data can be configured into appropriate data sets which will be analysed and

statistically tested.

Figure 10 Excel document master list

24

4.1.2 Data analysis

Once the data has been complied into a master list it is ready to be sorted into

relevant data sets such as species diversity by zoogeographical region. The Data

was extracted from the master list into a separate excel document which contained

country data base with zoogeographical region See (Figure 11). The species number

can be added from the master list by using the search tool for each country will give

a number of cells with that country name you can look through the cells selected for

errors the number then can be added to the new spreadsheet. Once all the data has

been placed in to the spreadsheet the data can be used to create graphs which

show species distribution between the zoogeographical zones.

Figure 11 Species richness by country

This spread sheet can also be used to create a heat map using online programs

such as Target map which was used in this study, although other computer

programs such as ArcGIS. As all the geographical data and species number are both

in the spreadsheet. The data can be uploaded to create a map to show spatial

relationships between species richness and political geographical regions. The

genus distribution has been calculated from the master list the genus column can be

ordered into alphabetical order from this the zoogeological data can be exported

from each species within the genus. This shows genus distribution this can be data

can be placed into a new spreadsheet to make data the analysis process less

complicated spreadsheet. This data can be used to create a pie chart to show

25

percentages of genus diversity by zoogeological region. As the Master list data has

been entered by hand typing there is potential for spelling errors which could affect

the data results so care must be taken in overlooking especially distribution data and

zoogeographic region as these are the independent variables whereas the species

names are not as important as species richness is being measured (number of

species).

4.2 Data statistical analysis

4.2.1 Relationship between endemic species and Zoogeographic

Region

The statistical tests carried out upon the data were the T-test to show significance of

endemic species in comparison to non-endemic species in each zoogeographic

region

4.2.2 Relationship between Genus richness and Zoogeographic

region

T- Test was carried out to test the significance of genus richness compared to

zoogeographic region. The data has to be carried out upon the mean of species

richness as the t-test requires two sets of data to compare against.

4.2.3 Relationship between Species richness and Zoogeographic

region

T- Test was carried out to test the significance of species richness compared to

zoogeographic region. The data has to be carried out upon the mean of species

richness as the t-test requires two sets of data to compare against.

4.2.4 Relationship between Genera richness and Species richness

in relation to zoogeographic region

Chi-squared was used to test the significance between genera richness and species

richness to zoogeographic region.

26

4.3 RESULTS AND STATISTICS

Figure 12 Pie Chart: World cockroach species richness by zoographical region

The pie chart in (Figure 12) Cockroach species presence shown by zoogeographic

region, based on 4549 species on the Cockroach Species File. the T-test P=0.46.

AUSTRALASIAN 9.41%

AUSTRO-ORIENTAL

11.96%

ETHIOPIAN 24.52%

MALAGASIAN 2.51%

NEARCTIC 1.05%

NEOTROPICAL 30.01%

NEW ZEALAND & POLY. 0.71%

ORIENTAL 7.92%

PALAEARCTIC 11.30%

CICRUMTROPICAL 0.26%

UNKOWN 0.34%

World cockroach Species Richness by zoographical region

27

Figure 13 Pie Chart World cockroach genera richness by zoographical region

The The pie chart in (Figure 13) shows Cockroach genera presence shown by

zoogeographic region, based on genera data on the Cockroach Species File. The T-

test P=0.5.

AUSTRALASIAN 8.03%

AUSTRO-ORIENTAL

13.67%

ETHIOPIAN 17.87%

MALAGASIAN 8.63% NEARCTIC

2.64%

NEOTROPICAL 22.66%

NEW ZEALAND & POLY. 4.68%

ORIENTAL 14.99%

PALAEARCTIC 6.83%

World cockroach Genera Richness by zoographical region

28

Newzealand &

poly

Australasian

Austro-oriental

Oriental Palaearctic EthiopianMalagasia

nNeotropic

alNearctic

Non endemic 36 13 76 100 50 11 3 25 24

Endemic 93 556 668 487 276 746 164 1345 26

0%

10%

20%

30%

40%

50%

60%

70%

80%

90%

100%

Spe

cie

s p

erc

en

tage

of

en

de

mic

ne

ss

Species endemic percentage by zoogeographic region

0%

5%

10%

15%

20%

25%

30%

35%

40%

45%P

ece

nta

ge o

f to

tal g

lob

al r

ich

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ss

World Cockroach Species and Genera percentage comparison by zoogeographic region

Species Richness

Genera Richness

Figure 14 Bar Graph World Cockroach species and genera percentage comparison by zoogeographic region

This graph shows the relationship between species richness and genus richness

based on genera data on the Cockroach Species File. The Chi square P=0.00

This Bar chart shows species endemism in each zoogeographic region as

percentages of the overall species richness in each area. The T-test provided a P=

0.005

Figure 15 Bar Chart: Species endemic percentage by zoogeographic region

29

Figure 16 Bar Chart: Biodiversity hotspots of Blattodea of species diversity

This Figure shows countries with over 100 species of cockroaches these 14

countries represent the highest areas of cockroach species richness in the world

from the Cockroach species file Data.

574

574

379

190

185

180

165

142

136

124

123

116

115

109

0 100 200 300 400 500 600 700

Australia

Brazil

Indonesia

China

South Africa

Democratic Republic of the Congo

Malaysia

Cameroon

India

Madagascar

Suriname

Panama

Colombia

Mexico

Biodiversity Hotspots of Blattodea of species diversity

Species Number

30

World

map

tha

t sh

ow

s c

ockro

ach s

pe

cie

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hn

ess b

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ach

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ivid

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ach

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. Cre

ate

d b

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sin

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are

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Figure 17 World map of cockroach species richness by political region

World

Map

of c

ockro

ach

sp

ecie

s ric

hn

ess b

y c

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31

5.0 Discussion

5.1 Percentage of Endemic species by zoogeographic region

This study showed a there was a significant difference in the percentage of endemic

species between regions (T-test P=0.005), thus the alternative hypothesis was

accepted and the null rejected. The highest percentage endemism (97% ≥) found in

Australasian, Ethiopian, Malagasy and Neotropical zoogeographic regions this

indicates these areas have the highest percentage of unique species in comparison

to the other regions. The high percentage of endemic species in each realm provides

further evidence that the zoogeographic regions are phylogenically distinct from

other regions (Holt et al, 2012).

The other regions Austro-oriental (89%), Oriental (82%) and Palearctic (84%) have

transitional zones where species overlap (Proches et al, 2012), New Zealand and

Polynesia have 74% endemic species. The reason for lower endemic species in this

region could be partly due to adventive species; cockroaches are renowned for their

survivability and could have got to the islands of Polynesia through various natural

pathways including surviving and travelling on drift wood or other plant material such

as coconuts. The other method is anthropogenic introduction, either intentionally or

inadvertently from travelling and surviving on shipping vessels or aircrafts (Vitousek,

1997).

This analysis of endemic richness can be used to help prioritise areas for future

conservation, when combined regions of high species richness this would assist in

identifying areas with high unique species (Keir et al, 2009). These regions can be

identified as areas which have a large contribution to Blattodea global biodiversity

such as the Neotropical region which boasts 98% (1345) endemic species.

The region where endemic species is lowest is in the Nearctic region with (52%

endemic species), which is considerably lower than any other region. However this

region has considerably lower species richness then the other regions. When this

lower percentage of endemics is combined with low species richness (50 species

and 26 endemic) it highlights the comparatively low diversity. This could suggest that

the Nearctic region is of a lower conservation importance of cockroaches

(Krishnamurthy, 2003). This comparison however does not take into consideration

32

higher phylogeny i.e. genera and family, if the 26 species belong to families unique

to the region it would increase their conservation status. This low endemic

percentage also suggests the border between Nearctic region and Neotropical

region is not in the correct location in the case of Blattodea as the transitional zone is

at too high level of crossover of species.

The (Figure 15) also provides evidence that the zoogeographic regions for terrestrial

invertebrates concept as endemic species percentages are very high in the majority

of the regions, although in some cases (New Zealand and Polynesia) can be

explained from adventive species or anthropogenic introductions.

5.2 World distribution map of species richness

The world distribution map (Figure 17) shows countries coloured by the number of

cockroach species present in each. The data was extracted from the Cockroach

Species File in 2014 and may not be a true representation of the world richness due

to species which have not yet been discovered or described. Mapping at the country

level using political boundaries has disadvantages, for example to cause a resolution

in Alaska being grouped with the remainder of America (with 50-80 species of

cockroach) this is not representative of Alaska. Spain is also another example as

Spain has many islands so mainland Spain cockroach species richness would differ

from the species found upon the islands. This also does not show geographical

barriers or distribution of the species within the country as a result of varied habitats.

However this gives a representation of species in each individual political region

rather than the actual distribution of the species. From the map countries with high

cockroach species can easily be identified such as Australia, Brazil and Indonesia as

these countries have 200 or more cockroach species. The countries with the most

species richness coincide with previously recognised biodiversity hotspot areas

primarily found within less economically developed countries (LEDCs). The map

also shows a disproportional species richness distribution as countries have

contrasting species richness, for example there is a higher majority of countries

above the Tropic of Cancer which have lower species richness (0-20 species) in

comparison to the equatorial zone especially in the southern hemisphere, with the

exception of parts of China and India.

33

Cockroaches are known to be found in tropical regions this map reinforces this know

perception as the countries with high species richness can be seen between the

Tropic of Capricorn and the Tropic of Cancer. There are the exceptions such as

Chad, Sudan and Angola however there could be a variety of reasons such as,

desert habitat is highly prevalent in these countries. The countries with low species

richness in Africa and the Middle East seem to correlate to the map of deserts in the

literature review Figure 7 (Hendrix et al, 2014). It has been noted that desert habitats

are less species diverse in comparison to other environments, as the Polyphagidae

family is the most prevalent in desert habitats in comparison to other families as

mentioned in the literature review (Edeny, 1974).

Other variables could be the lack of research carried out upon cockroaches, due in

part to the lack of taxonomists and funding within these countries. Europe and

Northern Asia and the Middle East are the areas where cockroach species richness

is lowest with the map showing 0-20 species in the vast majority of these areas.

These areas are dominated by temperate forest rather than tropical rainforest and

the low species richness is to be expected in these areas due to their minor

ecological role in temperate forests (Bell et al, 2007).

The counties with highlight in red in (200-574 species) Figure 17 areas have had the

most taxonomic research on Blattodea such as Australia by (Roth, 1991) China (Bey

–Bienko, 1970). These areas could have bias as there is a disproportional balance of

research and misrepresent countries with seemingly lower species richness

especially in the tropical regions. However without this invaluable research these

countries would possibly have considerably less known species and not be

highlighted with high species richness on the map. Also species bias is reduced in

the comparison of zoogeographic regions which is later discussed in more detail this

paper.

Other issues in which the raw data presented see appendix was countries such as

Yugoslavia no longer exist and these species have no specific locality on the

cockroach species file so cannot be reliably placed into 6 nations in which it has

become Bosnia- Hercegovina, Croatia, Macedonia, Montenegro, Serbia and

Slovenia (UN, 1996). This can become a problem with political regions overtime

they can change in size, split and change name for instance there are no Blattodea

34

records on the cockroach species file from South Sudan as it became the newest

nation in 2011 (UN, 2011). There are 21 species located in Sudan with a possible

proportion of those species distributed in the region in which South Sudan exists,

which is now not applicable of known cockroach species richness.

The map (Figure 17) also highlights countries with questionable species richness

which could indicate these countries need more research and exploration such as

Bangladesh which indicates it has 0-5 species of cockroach where as India has 120-

150 species. Honduras in Central America is another country which stands out from

the surrounding cockroach richness of 80-120 of Mexico and 50-80 Panama. The

Western Sahara region has no data at all and is shown as grey on the map, other

countries may have low data due to political, economic or social issues, countries

such as Yemen, Somalia and Syria these countries are currently very dangerous and

unstable due to political and economic issues so research is very difficult to conduct

in these specific regions (Collier, 2011). Identifying these regions could important for

the future and conservation cockroaches as these area can be revised and then

provide more data for analysis to improve the understanding and knowledge of

cockroach distribution and species presence.

Other consideration to take into account are habitat variation in each country for

example Gabon has lowland rainforest which covers 84% of the countries landmass

this is predominantly one habitat type (Butler, 2013), because of this the country

could have less species diversity in comparison to another with many different

habitat types such as Australia (Cousin et al, 2008). Other factors such as altitude

and climate cannot be seen as variables on species richness by country, also

historic data has been used so species which maybe locally extinct or extinct in the

wild maybe included and not truly represent global species richness however due to

data deficiency’s in Blattodea species population we currently do not know so are

presumed extant unless proven otherwise (Williams, 1996).

The map has its disadvantages previously mentioned; this map can be used as

reference point for global species richness as the data from the cockroach species

file which is a reliable source. The map shows the data currently known across the

world and therefore shows where there can be improvements and suggests areas for

research and focus conservation efforts for Blattodea. With only 24 species on the

35

IUCN redlist, this is a cause for concern as presumably all the other species are data

deficient (IUCN redlist, 2012).

5.3 Blattodea hotspots

The (Figure 16) shows countries with over 100 species of cockroach the highest

countries these should be focused upon for conservation of cockroaches as this is

where the majority of the world cockroach species are located and the ecosystem

services they provide is most likely to be highest (Bell et al, 2007).

The countries with the highest species richness are Brazil and Australia both

containing 574 species both of these countries cover very large areas, and contain

tropical rainforest and have a wide variety of habitat types. Also these countries are

more economically developed countries (MEDCs) and have provided funding and

facilities for taxonomists to research the same can be said for South Africa with 185

species. As previously mentioned the influence of taxonomists in specific countries

may introduce bias into the analyses (Sastre, 2009). The majority of countries are

less economically developed countries LEDCs according to the human development

reports (HDR, 2014) many of these countries have high deforestation rates with

Indonesia currently highest in the world with 840,000 hectares per year and

increasing deforested per year and Brazil second highest at 460,000 hectares per

year (Margono et al, 2014). This could potentially affect species richness in these

countries already however there is not enough data on the vast majority of species to

know whether they still are extant or not.

5.4 Species richness by Zoogeographic region

Understanding species richness is essential for conservation and understanding the

ecology of cockroaches, this gives a global overview of species richness by

zoogeographic region. (T-test P=0.46) thus the null hypothesis is accepted and the

alternative hypothesis was rejected.

However the species richness pie chart (Figure 12) shows how the cockroach

species are distributed around the world in each zoogeographic region. The

36

Neotropical region has the highest proportion of species; with 28.93% of the world

species are located in this region. This is to be expected as the Neotropical region

has the largest area of rainforest habitat in comparison to the other regions see map

in (Figure 12) in the literature review (Britannica, 1997). The Ethiopian region is

second highest with 15.98% and Austro-Oriental region 15.71% these regions both

have large areas of rainforest (Britannica, 1997). The regions with the lowest species

richness are Malagasy region with 3.53%, New Zealand and Polynesia 2.27% and

the lowest region Nearctic at 1.06%. These regions excluding the Nearctic are

mainly comprised of islands, islands are known for their low species richness

compared to mainland regions (Keir et al, 2009).

Species richness represented by zoogeographic region reduces bias from individual

countries with high taxonomic research excluding Australasian as it is only

comprised of Australia. As each country has species richness value an average can

be calculated over the entire region. Taxonomic bias occurs when taxonomists

regularly select areas with specific characteristics, this can affect the overall results

leading to a non-representative species richness of specific countries (Sastre, 2009).

The species with unknown distribution data are practically useless as there is no

locality data. Therefore they cannot be studied within the environment or

behaviourally and cannot be protected. However this is not a major issue as only

0.34% (21 species) of the known world Blattodea distribution data is unknown.

These species still have taxonomic importance as these species have been

described and catalogued. The lack of distribution information could have been down

to a variety of reasons such as poor curation practices; poor record keeping in the

field collecting or the data was just misplaced (Jahnke et al, 2012).

Phylogenetic variation and higher taxonomy level divergence such as families are

not taken in to consideration as only the species richness is measured. This must be

taken into consideration when comparing regions with species richness as this can

be misleading as the Neotropical region has the highest species richness. However,

when taking the phylogeny and higher taxonomy classes into account in the

Neotropical region which may have different conservation importance in comparison

to other regions.

37

5.5 Relationship between Genus richness and Zoogeographic

region

There was no significant difference in the richness of genera in comparison to

zoogeographic region. (T test P= 0.50) thus the null hypothesis is accepted and the

alternative hypothesis is rejected. The Pie chart in (Figure 13) represents genera

richness distribution by zoogeographic region; the regions with the highest proportion

of genera richness are the Neotropical with (30.20%) and Ethiopian region with

(24.67%).

The regions with the lowest percentage of global genera richness are New Zealand

and Polynesia (0.71%), Nearctic (1.06%) and Malagasy (2.52%). These regions all

have in common low species richness so is expected to be lower than the other

regions, also these regions are generally have less land mass and are made up of

predominantly islands excluding the Nearctic region. The other regions have similar

percentages Australasian (9.47%), Austro—Oriental (9.47%), Oriental (7.97%) and

Palearctic (11.37%).

When comparing genus richness often higher level relationships are no taken into

consideration as genera can vary in relation to one another, because they maybe in

different families. A Phylogenetic study by zoogeographic region would be a better

way of studying species divergence and would be able to provide greater evidence

of the zoogeographic theory in the terrestrial invertebrate’s field (Foottit et al, 2009).

5.6 Relationship between Genera and Species richness by

Zoogeographic region

There was a significant difference between genera and species richness in the

zoogeographic regions. (Chi squared P= 0.00) thus the alternative hypothesis is

accepted and the null hypothesis rejected. Comparing species richness to genera

richness by region can assist identifying regions which may be overlooked for their

importance to taxonomy. Species richness and Genera richness generally

correlates as there is low number of species in each genus (Adams, 2010).

The Neotropical region has both the highest percentage of genera and species

richness, however in relation to genera and species richness comparison overall is

38

noticeably higher on a global scale. This suggests that the Neotropical region may

not be as taxonomically important in comparison to other regions where genus

richness is greater than species richness. This is most obvious in the Malagasy and

the oriental region, where the proportion of genera is greater than the species

richness. This suggests these regions have higher phylogenetic diversity and also

could indicate these regions need more taxonomic research as there is a greater

potential for higher species undescribed or undocumented species.

5.8 How the study could be improved

The addition of Termitidae sister group data and a comparison on

distribution data would enhance the study this would improve the study from a

phylogenetic perspective by enabling the analysis of the relationship between

Blattidae and Termitidae distribution and zoogeological region (Eggleton, 2000). The

study of world termite distribution could be replicated however using the cockroach

data in order to make a meaningful comparison between the data two sets.

6.0 CONCLUSION

In conclusion the results showed there were significant differences in endemic

species in comparison to non-endemic species by zoogeographic region. This result

provided evidence for the zoogeographic region theory developed by Alfred Russel

Wallace however there was the anomaly of the Nearctic region, the results suggests

that the Nearctic region may not correctly placed for Blattodea. As there is high

percentage of non-endemic species as there is high transition between the Neotropic

region.

Overall the species richness in relation to zoogeographic region suggested there

was no significant difference this may be due to the vast amount of variables within

each region as these are not taken into account when comparing zoogeographic

regions. The data in which the analysis was based upon could possibly be improved

and therefore have an effect on the outcome of the statistical analysis. From the

various analyses of the zoogeographic differences these with notably Neotropical

region containing the most species. Regions with high species richness provide

39

evidence with the notion that cockroaches are predominantly a tropical species, the

regions with low tropical rainforest regions have lower species richness in

comparison shown in the Palearctic and Nearctic region. The evidence of species

richness corresponding cockroach world species richness map (Figure 16) with

habitat is shown with desert and tropical rainforest maps presented in the literature

review.

The future of cockroach species existence is unknown this is due to the lack of

conservation efforts for cockroaches is extremely poor with only two species with

data on the IUCN redlist (Gerlach, 2012). This most probably extends from the data

deficiency of species population and the overall disinterest of this order of insects.

More research has to be carried out for the data to be more reliable and accurate;

this would give a more up to date map of the species richness, as many of the

species from the current data set are presumed extant however many species could

be extinct locally or potentially globally extinct. The conservation of cockroaches is

dependent on this data to maintain future populations in a world which is changing

from a variety of anthropogenic effects such as global warming, deforestation and

pollution.

Brazil and Australia are pioneering examples of where taxonomic research has

produced vast amounts of detailed records with many of the records recorded to

regions within the countries. This level of taxonomic research was possible due to

the museum of natural history and research institutions and taxonomists specifically

interested in cockroaches. This has possibly caused country bias in cockroach

distribution, however the knowledge gained from this has been invaluable for

cockroach taxonomy and species richness.

However it must be taken into consideration that the data from the cockroach

species file is the currently known species distribution about the species may not

represent the actual global distribution. This is could be a significant issue as there

is a possible considerable difference in the knowledge of species distribution

currently then in the near future. The maps and graphs created in this study can be

used as a comparison in the future to see how the taxonomy and species richness

may have changed.

40

7.0 FURTHER RESEARCH

Countries which were highlighted in the world species richness map such as Bhutan,

Bangladesh, Cambodia and Honduras should be revised. As the species richness is

highly disproportional to the surrounding countries, there is potential for new species

to be described and to enhance the existing knowledge of species richness. This

would increase knowledge of species richness and create a more complete map in

the future in which the one in this study can be compared to see improvements of

taxonomic research.

The species richness map could be improved by using a different method such as

that used by Paul Williams on bumble bee distribution. He compiled a map of the

world into equal-area grid map excluding Antarctica with grid squares 611,000km

squared; this creates a map which is more specific for species richness to smaller

equal size areas rather than whole political regions which vary in size immensely.

This would be a better method when comparing areas as they are all the same size,

this still would not take into account habitat types however reduce the variables

which may affect species richness in each area (Williams, 1996).

Phylogenetic study in relation to zoogeographic regions would provide more

evidence to areas with higher taxonomic importance due to evolutionary uniqueness

and the clades in which the region contains. This data would drastically improve the

reliability of the study in order to prioritise areas for conservation and research efforts

(Holt et al, 2013). This could also provide data for studies such as the terrestrial

zoogeographic region update with invertebrate distribution data increasing the

accuracy of the zoogeographic regions to a wider range of taxa.

41

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9.0. APPENDICIES

9.0.1. Appendix 1

Country Name Region Species Number

Afghanistan PALAEARCTIC 6

Albania PALAEARCTIC 1

Algeria PALAEARCTIC 10

Andorra PALAEARCTIC 1

Angola ETHIOPIAN 15

Antigua and Barbuda NEOTROPICAL 2

Argentina NEOTROPICAL 30

Armenia PALAEARCTIC 0

Australia AUSTRALASIAN 574

Austria PALAEARCTIC 9

Azerbaijan PALAEARCTIC 4

Bahamas, The NEOTROPICAL 10

Bahrain PALAEARCTIC 0

49

Bangladesh ORIENTAL 0

Barbados NEOTROPICAL 4

Belarus PALAEARCTIC 0

Belgium PALAEARCTIC 4

Belize NEOTROPICAL 4

Benin ETHIOPIAN 8

Bhutan ORIENTAL 1

Bolivia NEOTROPICAL 38

Bosnia and Herzegovina PALAEARCTIC 1

Botswana ETHIOPIAN 26

Brazil NEOTROPICAL 574

Brunei AUSTRO-ORIENTAL 0

Bulgaria PALAEARCTIC 11

Burkina Faso ETHIOPIAN 0

Burma ORIENTAL 68

Burundi ETHIOPIAN 12

Cambodia ORIENTAL 16

Cameroon ETHIOPIAN 142

Canada NEARCTIC 1

Cape Verde ETHIOPIAN 5

Central African Republic ETHIOPIAN 7

Chad ETHIOPIAN 8

Chile NEOTROPICAL 14

China ORIENTAL

China PALAEARCTIC 190

Colombia NEOTROPICAL 115

Comoros MALAGASIAN 8

Costa Rica NEOTROPICAL 69

Côte d'Ivoire ETHIOPIAN 4

Croatia PALAEARCTIC 2

Cuba NEOTROPICAL 60

Cyprus PALAEARCTIC 3

50

Czech Republic PALAEARCTIC 7

Democratic Republic of the Congo ETHIOPIAN 180

Denmark PALAEARCTIC 3

Djibouti ETHIOPIAN 0

Dominica NEOTROPICAL 9

Dominican Republic NEOTROPICAL 15

Ecuador NEOTROPICAL 67

Egypt PALAEARCTIC 11

El Salvador NEOTROPICAL 0

Equatorial Guinea ETHIOPIAN 32

Eritrea ETHIOPIAN 6

Estonia PALAEARCTIC 0

Ethiopia ETHIOPIAN 59

Federated States of Micronesia NEW ZEALAND and POLY. 0

Fiji NEW ZEALAND and POLY. 5

Finland PALAEARCTIC 2

France PALAEARCTIC 13

Gabon ETHIOPIAN 29

Gambia, The ETHIOPIAN 3

Georgia PALAEARCTIC 6

Germany PALAEARCTIC 8

Ghana ETHIOPIAN 25

Greece PALAEARCTIC 14

Grenada NEOTROPICAL 10

Guatemala NEOTROPICAL 51

Guinea ETHIOPIAN 66

Guinea-Bissau ETHIOPIAN 5

Guyana NEOTROPICAL 59

Haiti NEOTROPICAL 23

Holy See PALAEARCTIC 0

Honduras NEOTROPICAL 11

Hungary PALAEARCTIC 11

51

Iceland PALAEARCTIC 0

India ORIENTAL 136

Indonesia AUSTRO-ORIENTAL 379

Iran PALAEARCTIC 18

Iraq PALAEARCTIC 8

Ireland PALAEARCTIC 0

Israel PALAEARCTIC 1

Italy PALAEARCTIC 36

Jamaica NEOTROPICAL 35

Japan PALAEARCTIC 34

Jordan PALAEARCTIC 1

Kazakhstan PALAEARCTIC 6

Kenya ETHIOPIAN 74

Kiribati NEW ZEALAND and POLY. 0

Kuwait PALAEARCTIC 0

Kyrgyzstan PALAEARCTIC 8

Laos ORIENTAL 8

Latvia PALAEARCTIC 1

Lebanon PALAEARCTIC 0

Lesotho ETHIOPIAN 17

Liberia ETHIOPIAN 8

Libya PALAEARCTIC 8

Liechtenstein PALAEARCTIC 0

Lithuania PALAEARCTIC 0

Luxembourg PALAEARCTIC 2

Madagascar MALAGASIAN 124

Malawi ETHIOPIAN 29

Malaysia AUSTRO-ORIENTAL 165

Maldives ORIENTAL 0

Mali ETHIOPIAN 5

Malta PALAEARCTIC 0

Marshall Islands NEW ZEALAND and POLY. 0

52

Mauritania ETHIOPIAN 2

Mauritius MALAGASIAN 0

Mexico NEOTROPICAL 109

Moldova PALAEARCTIC 2

Monaco PALAEARCTIC 0

Mongolia PALAEARCTIC 1

Morocco PALAEARCTIC 38

Mozambique ETHIOPIAN 70

Namibia ETHIOPIAN 43

Nauru NEW ZEALAND and POLY. 0

Nepal ORIENTAL 11

Netherlands PALAEARCTIC 1

New Zealand NEW ZEALAND and POLY. 23

Nicaragua NEOTROPICAL 26

Niger ETHIOPIAN 23

Nigeria ETHIOPIAN 21

North Korea PALAEARCTIC 1

Norway PALAEARCTIC 1

Oman ETHIOPIAN 0

Pakistan ORIENTAL 2

Palau NEW ZEALAND and POLY. 1

Panama NEOTROPICAL 116

Papua New Guinea AUSTRO-ORIENTAL 68

Paraguay NEOTROPICAL 26

Peru NEOTROPICAL 92

Philippines AUSTRO-ORIENTAL 88

Poland PALAEARCTIC 4

Portugal PALAEARCTIC 14

Qatar PALAEARCTIC 0

Republic of the Congo ETHIOPIAN 18

Romania PALAEARCTIC 11

Russia PALAEARCTIC 6

53

Rwanda ETHIOPIAN 27

Saint Kitts and Nevis NEOTROPICAL 1

Saint Lucia NEOTROPICAL 3

Saint Vincent and the Grenadines NEOTROPICAL 10

Samoa NEW ZEALAND and POLY. 11

San Marino PALAEARCTIC 0

Sao Tome and Principe ETHIOPIAN 0

Saudi Arabia PALAEARCTIC 4

Senegal ETHIOPIAN 14

Seychelles MALAGASIAN 21

Sierra Leone ETHIOPIAN 13

Singapore AUSTRO-ORIENTAL 22

Slovakia PALAEARCTIC 3

Slovenia PALAEARCTIC 0

Solomon Islands AUSTRO-ORIENTAL 7

Somalia ETHIOPIAN 17

South Africa ETHIOPIAN 185

South Korea PALAEARCTIC 2

Spain PALAEARCTIC 64

Sri Lanka ORIENTAL 51

Sudan ETHIOPIAN 21

Suriname NEOTROPICAL 123

Swaziland ETHIOPIAN 21

Sweden PALAEARCTIC 0

Switzerland PALAEARCTIC 11

Syria PALAEARCTIC 10

Taiwan ORIENTAL 54

Tajikistan PALAEARCTIC 12

Tanzania ETHIOPIAN 96

Thailand ORIENTAL 41

The Former Yugoslav Republic of Macedonia PALAEARCTIC 1

Togo ETHIOPIAN 22

54

Tonga NEW ZEALAND and POLY. 1

Trinidad and Tobago NEOTROPICAL 40

Tunisia PALAEARCTIC 13

Turkey PALAEARCTIC 16

Turkmenistan PALAEARCTIC 4

Tuvalu NEW ZEALAND and POLY. 0

Uganda ETHIOPIAN 32

Ukraine PALAEARCTIC 4

United Arab Emirates ETHIOPIAN 0

United Kingdom PALAEARCTIC 3

United States of America NEARCTIC 63

Uruguay NEOTROPICAL 6

Uzbekistan PALAEARCTIC 7

Vanuatu NEW ZEALAND and POLY. 2

Venezuela NEOTROPICAL 78

Vietnam ORIENTAL 95

Yemen ETHIOPIAN 3

Yugoslavia PALAEARCTIC 16

Zambia ETHIOPIAN 36

Zimbabwe ETHIOPIAN 66

9.0.2 Appendix 2

See CD-ROM attached contains the Master world Cockroach species file