SECTION A THE BACKGROUND

PRUT ACS’.

Any

serious

stuck

ot

the Biosystematics

and

Ecophysiology

of a group of organisms will constantly face the problem of the origin and

evolution

oi

the

ecosystem

and

their abiding

flora and fauna.

The origin and evolution of the fauna of a particular region is closely linked with the origin and evolution of the flora and the coevolution of both is directly correlated with and profoundly influenced by the biogeomorphological

evolution

of

the

region.

The

biogeographical

and geomorphologicai evolution could be better understood only when they are studied in relation to the historical background of the decisive changes that A

have taken place in the

meaningful interpretation

of

any

land mass in space and time. ecophystological

component

of

an organism is possible only when the biogeography of the particular region is clearly understood.

It region ^

is

an

like

established

everything

fact that

else,

the climate of

constantly

changes.

a particular

The

prevailing

climatic conditions of a given region have not given rise to the pre­ sent day pattern of distribution ol

both the flora and the fauna and

the ecophysiofogical attributes of the flora and fauna is nothing but an expression of the interaction between the organism and physico­ chemical elements of the inicrohabitat. Both

cannot

be

studied

in

isolation.

the

According

to

Mani

Biogeography

of

the

continues

to

influence

and

(1974), Indian

tire

most

Subcontinent

determine

remarkable is

the

feature of

Himalaya

that

the climate of the whole of

2

India.

The

uplift

of

the

Himalaya

Pleistocene, in conjunction with late

Tertiary,

affected

lias altered

in

the

late Pliocene and early

the tiplilt of the Western Ghats in

the pattern of rainfall, and concurrently

the pattern of distribution ol

both fauna and flora. It is

a well established fact that the entire Peninsular fauna was originally of a I tumid Tropical Rainforest one. which was far more extensively and continuously distributed throughout and according to

the Peninsula than at present

Vlaru 11974) changes in the habitats affecting the

distribution have taken place within historic times. The present day climate of India does not underlie the origin of biogeographical charac­ ters.

Instead,

present

day

they

are

distribution

products ol therefore

geomorphoiogical evolution. The represents

a

dynamic

phase

ot

uninterrupted course of biogeographical evolution that has no means neither stoppednor even substantially slowed down.

According to Mehr Homji (1974), the continental drift, that caused the uplift of Himalaya, is directly and indirectly responsible for the climatic changes, directly by pushing Southern India northward from its closeness to the equatorial line and indirectly by the lifting of the lofty Himalayan system, that continue to influence the com position and ecophysiology of the flora and fauna. The monsoon is defined as

year having 100 rainy days and the rainfall in the summer

reaching the maximum. According to Mehr Homji (1974), this definition of

monsoon

is

appropriately

applicable to Southern India in which

the annual rainfall ranges from 1000 to 1500 mm and the average duration of dry ^.season is about six months and the mean temperature of the coldest month not below 20°C. In the summer months (May -

3

June), the rainfall ranges from 2>i i. 200

mm

36'f mm. If the rainfall exceeds

it is the region of wove! humid climate, favouring tropical

wet ever green forest and when the summer rainfall • is less than 200 mm and irregular in May and Dune

it signifies the domain of

the

tropical dry evergreen forest or the Scrub Jungles. When the rainfall is below 100 mm it characterises rim domain of the Semiarid zone. Both Semiarid /.one and Scrub Jungles completely and the Agroeco­ system partly could be det lared as drought prone ecosystems.

Through

destruction

agencies within historical

ol

retinal

habitats,

induced

by

human

times^ extensive telling of trees accelerated

the deterioration of powers of adjustments in the ecosystem. Rapid industrialisation witnessed

a

as

a

concomitant

progressive

process

deterioration

of

of

modern civilisation,

climate,

elimination

and

destruction of habitats, impoverishment of flora and fauna, regression ♦

of distributional ranges and creation of ecological islands.

Consequently, such biogeographical evolution lead to unstable conditions of microhabitats and to the evolution of isolating mechanisms that These

promote

formation

of

ecotypes,

biotypes

and

created so many other complicated problems

taxonomist.

The

sibling

species.

for a museum

insects, constituting the major participants in

the

changes of an ecosystem^ could be considered to be most successful among all other animals and one of the major components of such, a dynamic class is the Reduviidae, popularly known as the 'Assassin bugs'. Intensive Bharathiar

survey

University,

conducted located

in

by the

the

research team

Paighat

Gap of the

of

the

Western

4

Ghats, lor over a decade. has < un< S lovely established the fact that the to

Reduviidac of 6r> genera and

this 1 1

region, comprising

170 species belonging

subfamilies, constitutes an army of predators

of insect pests_, as well as millipedes of the diverse ecosystems of this

region.

Proper

understanding o|

ecosystem will pay divideids, il harnessed.

Out

the

management of the

Agro

these natural predators are discretely

of the about 40

species described new to science,

10 spec ies belong to ibe nontibieroi mie group and more than 60 species are new records, not listed eariiei

bv any of the workers, from this

region. The thesis presented here summarises the investigations carried out in the Division of Entomology, Bharathiar University, for a period of four years. Frequent field trips were made by the team of workers in this D.S.T. funded project.

Most of the collection spots are in•

tested

with

venomous

reptiles

and

scorpions

making

collections

a

very risky endeavour. Light trap collections were made to investigate species abundance during different seasons. Considering the enormity of the work undertaken and the magnitude of the materials accumu­ lated, 1 he author has to bo satisiied in presenting the informations on the biosystematics and ecophysiology of. only the nontibiaroliate group of Reduviidae that constitutes roughly 56% of all the reduviid fauna accounted for from this region.

The thesis begins with a brief taxonomy and Biosystematics of

review of literature of the

the Reduviidae of the world, with

special reference to the Oriental Reduviidae. In the opening chapter

on materials and methods, places ol collection and the species collected have

been

serialised

methodology ol

systematically

study ol

the

and

plotted on

the maps.

The

lostrum, stylets, origin of tibiarolium,

parameres, pygophore spines, basal plates and the aedeagus and their modifications have been assessed for systematics. In the next section on Taxonomy, 95 species belonging to 39 genera so far recorded under this

group

from this region have been catalogued. One genus

and

18 species have been described new to science and a number of species have been reported as new records from Southern India.

In the ensuing section on Ecophysiology, the functional and evolutionary importance of the feeding strategies and the male genitalia have been assessed. Finally, an attempt has been made to interpret the structural features in relation to the origin and evolution of the non-tibiaroliate group of Rcduviidae from a Tropical Rainforest eco­ system to a drought prone ecosystem in Southern India. A brief summary is followed by a serialised list of over 350 alphabetically arranged references

have been cited

in the text. The thesis

is substantially supported by 7 maps, 41 text figures, 3 tables 52 3 photomicrographs, arranged in

21

and

plates.

It is the fervent hope that this discourse will provide yet another deck for a better interpretation of the origin and evolution of Reduviidae of Western Ghats.

BRIEF

REVIEW

OF

LITERATURE

The family Reduviidue < oukl i; .

< onsidered as one of the best

known families of Heteroptera. Eversince the erection of this group of predaceous bugs by Latreille (18-0A), a steady flow of information on the taxonomy of these bugs is inuicatcd by the accumulation of literature and ever since W igglesworth (I 9 32) picked up the haernophagous reduviid bug

Rhodnius prohxws as

was shifted

his

from taxonomy (Museum

laboratory

pet, concentration

taxonomy) to lab oriented in­

vestigations. At present a casual glance of the burgeoning literature on

Reduviidae

will

instantly

on the various aspects oi

reveal

a predominance of publications

Rhodnius proiixus alone and it is almost

a JUvTYmcru?) task to review the literature pertaining to this particular species alone. The reduviid next of interest, which has assumed greater importance due to the potentiality of its vector role is yet another haernophagous bug and

1953,

Luccna

1973, Forattini,

Tnatoma infestans (Baptist 1956,

1957,

1958,

1959,

1960,

1975, Sankaran and Nagarajan,

1941, Barth 1952 1961,

1965a and

1975 and Houseman,

1983). A reduviid taxonomist cannot proceed witfciout the guidance of

the significant

taxonomic

papers

published by

Usinger (1943 Sc

1966), Carayon (1944, 1954. 1958 Sc 1966), Carayon and Villiers (1968), Wygodzinsky

(1947,

1972.

197 5,

197 3,

1968, 1972, taxonomy Mexico;

of

1966

&

1980),

1976a A b and

and

Villiers (1968,

1969,

1971,

1986), Maldonado Capriles (1966,

1976a,b,c & d, 1986a, b & c and 1987a,b & c) on the the

Reduviidae

Maldonado capriles

of and

Venezuela,

Jamaica,

Belgium

Brailovsky (1983a & b and

and 1985)

7

on the Reduviidae of

Mexico; Har

some species of Reduvudae

from

have contributed

to

immensely

extra oriental species ol

(5972, 1975, 1986 and 1987) on America, Canada and West Indies knowledge on the taxonorm

>ui

of

Reduviidae. Wygodzinsky (1966) is singularly

responsible for our present knowledge of Emesinae and his monograph is a treatise on this subfamily.

Apart

from

these,

a

number

of

occasional

publications

on

isolated groups on Reduviidae have added further to the bulk of reduviid literature and most of these publications are on regional basis, parti­ cularly of the North American Reduviidae. These include the contri­ bution of

Readio (1927), Davis (1955,

Wygodzinsky (1947,

1963,

1966 and

1957,

1974),

1961,

1966 and

1969),

McMahan (1982 &

1983)

and Giacchi (1969, 1970, 1973, 1974, 1977, 1980, 1982, 1983, 1984 & 1985)

on

several

genera

of

American

Stenopodainae

Giacchi

und

Maldonado capriles (1983), Odhiambo (1958a, 1958b & 1959) and Villiers 1972 and 1973) have contributed on African Reduviidae. Other more significant publications on extra Oriential reduviid taxonomy include those

of

Wygodzinsky

Cryptophysoderes from

and the

Maldonado Capriles (1972) on the genus New

World,

Hsiao (1973,

1976a,

1976b

and 1977) on the Reduviidae of China, Else et ad. (1977) on a new species of

cave inhabiting

Pelaez (1981)

on

Triatoma from

the Salyavatinae

Maldonado Capriles (1983)

on

new

Malaysia, Brailovsky and

of North America, Giacchi and species of Stenopodainae from

Nicaragua and Malipatil (1983) on a new genus of Holoptilinae from Australia. Zarate and Zarate (1985) have given a check list of the Triatominae of Mexico. Putshkov (1981, 1982a, 1982b and 1983), Putshkov and Putshkov (1984), and Putshkov et al. (1987) further contributed

8

immensely to our knowledge on the Taxonomy of Russian Reduviidae. Dougherty (1982 &

1989) reviewed

the South American genus Z;£ta

with three new species and ci&scYib&l

ck

more genus of Ectrichodiinae

from South America.

The to the

contributions oi

families and

IVroursey

subfamilies of

(1963

&

1971) _ on

the key

the nymphs of North American

Hemiptera and China et al. (1939) on the check list and key to the families and subfamilies of the Hemiptera are considered outstanding.

The

taxonomy

ol

Oriental

Reduviidae

had

its

major

break

through when Distant (1904 &c 1910) made this significant contribution on the reduviid fauna of British India including Ceylon and Burma. He described 342 species belonging to 106 genera and 13 subfamilies and since then very few publications on Oriental species of Reduviidae have been documented. A.

Cook (1977)

made an attempt on

key for the identification of genera and species of Oriental Ectri­

chodiinae

based on

museums.

Apart

the collections deposited

from

this,

the

taxonomic

in

Indian and British

contributions

of

Indian

Reduviidae are limited to the descriptions of two species of Harpactorinae

by

Samuel

and

Joseph

(1953).

Subsequently,

Muraleedharan

A/

(1976) described two new species of Henricoha^ia from India, Ghauri (1976) described two more species of Linshcosteus and Bhargava arid Prasad (1977) a few more species of Reduviidae/V&m. Northwest India. Wygodzinsky and Lent (1980) added two more genera of Reduviidae and Ambrose and Livingstone (1986a and 1986b) added one new species of Harpactorinae and one new genus of Ectrichodiinae from Southern

9

India. Other recent publications on the Reduviidae of peninsular region include

those

of

Livingstone

and

Murugan

(1987).

Livingstone

and

Ravichandran (1988) who added a dozen more species to the list of Reduviids from Southern India.

Readio (1927 &

1931), Miller (1 953 & 1956), Odhiambo (1959)

and Decoursey (1963) are

mainly

responsible for

our knowledge on

the biology of Reduviidae. Subsequently, Wallace (1953), Gillett (1935 & 1957),

Parker

(1961,

1961a,

1963 6c

1965a,

! 965b.

1969

6c

1971),

Edwards (1960,

1966), Nyiira (1 970), Louis (1973) and Swadener and

Yonke (1973a, b & c), Habib (1977), Zeledon et_ ah (1970 & 197 3) and Shepard

et_ ah (1982) have contributed further to our present

information on the biology of a miscellany of extra Oriental reduviids. The biologies of Oriental reduviids are known only through the contri­ butions of Joseph (1959) on Rhinocoris lapidicola and JR. nysiiphagOs, Bose (1949), on Coranus spiniscutis, Livingstone and Ambrose (1978) on

Acanthaspis pedestris

Ambrose

and

Livingstone

(1979 6c

1985b)

on Lophocephala guerinl and Coranus vitellinus and Ambrose et al. (1985a) on Catamiarus brevipennis.

Regarding the ethological studies of Reduviidae the nutritional ethology

has received greater

Edwards, (1961, exsanguis,

1962 &

aAtam/fcum

since the publications of

1966) who initiated such studies on Zelus

Rhinocoris carmelita and Platymeris rhadamanthus. Subse­

quently Arnold (1971), Parker (1961, 1965, 1969, 1971 6c 1972), Zeledon et ah (1970 & 1973), Swadener and Yonke (1973), Stoner et ah (1975), Friend

and

Smith (1975),

Cobben

(1978),

McMahan (1982 6c

19831,

10

Eads and Campos (1983), published

scries of papers on the feeding

strategies of a few species of extra^oriental Reduviidae. The feeding strategies ol Onential reduvnds have been sttAdUexl and

Ambrose (1978a,

1978b

&

(1981), Ambrose et_ al. (1983,

1984), Haridass and 1989,

by Livingstone Ananthakrishnan

1985b and 1986a) and Venmson

and Ambrose (1987). By their investigation they were able to prepare models on the feeding strategies of both entomophagous, myriophagous and haemophagous reduviids of this region.

Apart

from

the

feeding

behaviour

attempts were made to understand their (1965b, 1969,

1971

&

of

Reduviidae,

very

few

mating behaviour. Parker's

1972) works on various species of Reduviidae

are considered to be pioneering in this regard. Ambrose and Livingstone (1978b,

1979b, 1984, 1 985a,

(1987) and

1985c and

1987), Ambrose and Vennison

Ambrose et_ aL (1986b) have highlighted certain

aspects

of the mating behaviour of some species of Reduviidae of Southern India. The

spermatophores

of

Reduviidae

to Rhodnius have evinced considerable

with

special

reference

interest after the works of

Davey (1959, 1960 & 1967) and Khalifa (1950a and 1950b) and in this connection the male internal organs of reproduction also have received considerable importance as revealed by the publications of Woodward, (1950), Davis, (1955), Kumar, (1964 6c

1967), and Louis and Kumar

(1973).

Presence of tibiarolium (tibia! pad' or fossula spongiosa' 'Spongy

furrow')

in

Reduviidae

or

has been considered as a structural

adaptation for capturing their prey (Distant 1904; MacGillivray 1923;

Poulton, Miller,

1931; Gillett 1938,

and

1939 and

Wiggle ,worth,

1942; Edwards.

1932;

I960,

Wigglesworth,

1961b,

1963 and

1938; 1966

Edwards and Tarkanian, 1970 . subsequently, Livingstone and Ambrose (1978a, 1978b and 1984) made an attempt on the evolution of 'tibial pads'

in relation to the predatory elliciency in a large number of

species of Reduviidae of the Western Ghats.

Detailed studies of the stylets and their functions during feeding act have been taken up by various workers. Most of their contributions were mainly on the kaernophagous bugs and

a

few

other

heteropteran

this regard include those of

Rhodnius and Triatoma

species.

Significant contributions

in

Butt (1943), Kullenberg (1946), Leston

(1957), Haskell (1 957), Lavoipierrc (1959), Banerji (1960), Pinet (1963, 1964 &

1968), Matsuda (1965), Friend (1965), Forbes (1966 & 1969),

Livingstone (1969), Bernard et_ al. (1970), Friend et_ aL (1971 & 1977), Pendergrast singularly

(1976)

and

responsible

Bowling (1979 &

for

our

present

1980). Cobben (1978)

is

knowledge on the stylets of

other entomophagous insects.

The stridulatory mechanism of some terrestrial species of Hemiptera

including

Reduviidae

has

been

worked

out

by

Miller (1 953),

Leston (1957), Haskell (1957), Schofield (1977) and Diluciano (1981). Most of them have focussed their attention on the stridulatory mechanism of some species of Triatominae.

Detailed studies on the digestive system, salivary glands and

tvsuM phylogenetic relationships among several reduviids have been reported by several workers such as Edwards (1961), Kumar (1962), Miles (1972),

12

Louis and

Kumar ( 1 97 3), Cobber \ 9/8) and Haridass and Anantha-

krishnan (1 980).

Regarding Brindley'seem chemistry

ot

secretion

glands

iiave been

LkeLn

structure,

nvestigated in

and

bio­

Rhodnius prolixus

and Triatoma infestans by Games et ah (1974) and Kalin and Barett (198 3). The contribution of I’atienddi m ah (1 972). Staddon (1974), Schofield (1977). are remarkable on islanding.

Haridass

and

Anunthukrishnun

(1981)

worked

on

the

pylorus and rectal glands of a wide range of Reduviidae of Southern India. Control of pests by

the Assassin bugs have been investigated

by various workers. Vanderplant (1958) reported Pfatymeris rhadamanthus predating

upon

the

Orvctes rhinoceros, upon cotton

boll

Rhinoceros beetles such as Oryctes boas and Morgan (1967) on

weevil,

Apiomerus spissipes predating

Vlukerjee (1971) on

Reduvius

sp.

feeding

on Corcyra c ephalonica, and recently Sitaramiah et_ ah, (1974 & 1976), Code et_ ai_. (1978) and .-Ambrose (1985) reported a few more species feeding

on

Heliothis

armigera,

Calocoris a ugustatus,

Earias

fabia,

Harvester^nts and Spodoptera litura.

Nymphal Camouflaging of Reduviidae have been thoroughly investi­ gated by Odhiambo (1958), Zeledon et_ al_. (1969 and 1973), Ambrose (1986) and Livingstone and Ambrose (1986) in Acanthaspidinae, Triatominae and Salyavatinae.

The population dynamics oi three species of reduviids in different habitats

has

been

worked

out

by

Ambrose and Livingstone (1978)

13

and

the

lunar

periodicity

and

light

trap

collections of

Reduviidae

and other heteropteran specie's were highlighted by Goel (1976) Onder (1979) and Kaushai and Vats (1984).

Pruthi (1935), Baker (1931). Gtli.-ii (1935), Elson (1937), Crampton (1938), Snodgrass (1935 and

1936),

Michener (1944), Carayon (1944),

Ashlock (1 957), Gillett (1957), Mar Ciliivray (1923a and 1923b), Dupuis (1955 and 1970), Matsuda (1958 and 1976), Scudder (1959), McDonald (1961,

1963 and

1982), Schaeler (1968 and

1977) and Cobben and

Wygodzinsky (1975) are mainly responsible for their present knowledge on the male and female genitalia of all Heteroptera including Rhodnius (V

prolixus and^few other Triatoma species.

The reference (1957,

comparative

morphology

to

was

1961,

genitalia 1966,

very

1969 &

of

the

Reduviidae, with special

elaborately

1975) and

worked

out

by

Davis

the contributions of Carayon

(1944 and 1968), Carayon et ah (1958 and 1968), Kumar (1962) and Cobben

and

Wygodzinsky

(197 5)

on

the

morphological

features

of

a number of reduviids are of great value to a reduviid systematise Subsequent

publications on

the

genitalia of

Reduviidae which have

great taxonomic value are from Lent and Jurberg (1966, 1967, 1969, 1972, 1983 and 1984) and Giacchi (1969, 1973, 1974, 1977, 1978, 1979, 1980,

1983,

1984 and 1985). The internal anatomy of a variety of

Reduviidae was

SUretied.

by Kumar (1962b) with special reference

to their reproductive system.

The foregoing review of literature pertinent to reduviid, morphology taxonomy and biology thus indicates that intensive studies of structural features and their taxonomic interpretations of these bugs omYegional A*

basis will go a long way

in

furthering the dimensions of reduviid

taxonomy and systematics in India.

14

MATERIALS

An intensive survey

AND

tor

METHODS

recording the reduviid fauna of the

states ol

Southern India has been tarried out during a period of three

years by

visiting

scattered

ail over the lour main

frequently vu ions collection

sites

ecosystemsof this region.

A total

of 170 species belonging to 66 genera and 11 subfamilies of Reduviidae have been so belonging

to

identified 19

iar recorded and cm:

species

conveniently

38 genera and

o!

these a total of 9b species

six subfamilies

have been

belong to the non tibiaroliate group. Out of these have in

been

described

new

seven geographical

to science.

maps (Maps

They are plotted

1-7).

numbers of species are given in the maps. The numbers given in parentiUmbefti

thesis represent the numbers found m the museum catalogue^maintained in the Division of Entomology. Bharathiar University. The meteorological data as well as the elevation collection sites ond. 're.ccrrzild

Xri

efc aM

the lour ecosystems were

for every collection trip using the 'Barigo' thermohygro-

qraph and Altimeter. Coimbatore district was most intensively surveyed run mtHjf

and seven sites representing mamlv Scrub Jungles and

Agroerosvstem

three ecosystems, Semiaridzones, were

selected for fortnightly re­

cording ol the population oi the life stages, sex ratio, etc. From these data as well as the general coli c ! ion data the spatial and vertical distribution ol all these species haw been tabulated and represented in the pie charts and histograms.

Most ol Rainforests.

All

the non tibiurolmte species are endemic to Tropical the spe. ies ot

im

genus Rhuphidosoma, the female

S.No.

M.C.No. Sub^fa mi!y

Slenopoda ina e

71.

(145)

Oncocejihalus cinga’cnsis U m‘(cr

72.

(146)

Oncocephalus klugi Distant

73.

(147)

Oncocephalus chanmndaii s/> . no\

74.

(148)

Oneoceplxa 1 us impi a 1 icus Re i;,' cr

75.

(149)

Oncocephalus morosus Disia;

76.

(150)

Oncocephalus annul iocs Slai

77.

(151)

Oncocephalus notalns l\lu •

78

(152)

Oncocephalus modestus Ren', ( 1 x

79.

(153)

• Oncocephalus schioedtei Reu ter

80.

(154)

Oncocephalus aterrimus !'>■■ ■■ • n!

81.

(155)

Oncocephaltis yashrxtiii s/n cr

82.

(156)

Oncocephalus bipun claim so. no\

83.

(157)

Caunus farina tor Reuter

84.

(158)

Diaditus errvbundus Distant

85.

(159)

Staccia diluta Slat

86.

(160)

Bardesanes sericonotatus s,"- now

87.

(161)

Sastrapada baerensprungi set

88.

(162)

Sastrapada dongata Sp. tin- .

89.

(163)

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