Byomya (all synonyms of tempestiva Fallen), and 15 species of Musca (all

synonyms

of autumnalis

and

domestica, exceptfor

one

subsequentlytrans- ferredtoPollenia(Calliphoridae)).

His system was

largelyfollowed

by Lioy

(1864),

who produced a

synthetic

work,

withoutoriginality

and

apparently inignorance

of most

ofthe

work of

his

fellow-countryman Rondani.

Lioy's

genera {Musca, Plaxemya, Byomya) and

theirdiagnoses

were

takendirectly

from Robineau-Desvoidy's work. When asked

for his verdict

on

Lioy's

work

(andhisreasonforignoringit),

Rondani

replied that "c'est

de

lapoe'sie.

"

Following from

Schiner, the Austrians F.

M. Brauer and

J. E.

von Bergenstamm

dealt

comprehensively

with the

European

Tachinoidea, to- getherwith

a

large

number of

exoticforms,

and

their

work

also

covered

the

groups of Muscidae

with

a

forwardly-curved vein

M.

^Theirinterpretationof

Musca and

allied

genera was

naturally

based on

that

of Rondani and

Schiner

and

theyaccepted

Rondani'

s separation

of Musca

^into

two

genera,

though any

meritinthisdivision

was

obfuscated

by

inaccuraciesintheirdifferentia- tion(Brauer

& Bergenstamm,

1889: 156):

Eyesbare,almost touchinginc?,separatedin$^,etc.

MuscaL.s.str.Rdi.,domestica L.

Eyeshairedinct,pubescentin$^.Fronsof^6*moderately broad, verybroadin9,etc.

PlacomyiaR.D.emend. {PlaxemyaR.D.)vitripennisMg.

This

distinction

was maintained

to the

end of

their

work

(Brauer

&

Bergenstamm,

1893: 179)

and

theauthorsalsoincluded

a check

listofthe species thatthey studied(Brauer

& Bergenstamm,

1891: 419),

which was

useful insofar as italso indicated

some

of theearly type/syntype material locatedinthe

Vienna Museum.

The

lastgreatsystematic

work of

thecentury

was by

the

Frenchman

L.

Pandell6

(1824-1905) and comprised a monograph of

the

"Muscidae"

(i.e.

both

calyptrates

and

acalyptrates)

of

France.

He

introduced

a number of new

characters

from

thechaetotaxy

of

the

body,

wings,

and

legs intothe

taxonomy of

the

Schizophora and

alsopioneered the study

of male

genitalia. Inde- pendently

of

hiscontemporaries,

he

discovered the value

of

thechaetotaxy

of

the

body and

legs, but the prolixity

of

his text

and absence

of

any

illustrations deprived

him of any

justly-earned credit for these findings.

Pandell6 (1898a: 20-23)

gave a key

tothe

genera of

his division"Musce'es"

that

was

quite different

from

that

of

his

immediate

predecessors, using

a

wealth

of

characters that

were mostly new. His

"Musce'es"included

Mesem-

brina, Pyrellia (in

a

very

expanded

sense),

Musca,

Stotnoxys,

Gymnodia, Graphomya

,

and Muscina. His genus Musca

(Pandell6,1898b: 40)isentirely

modern

in its

scope and

with

many

features inthegenericdescription that

26 Systematic Database of Musca Names

were

independently rediscovered

by

the next generation of taxonomists.

Four

species

were

included:vitripennis,tempestiva, corvina,

and

domestica.

Pandelie (1894, 1896)

and

Girscnner (1894, 1896, 1897)

were both

influenced

by

the

work on

chaetotaxy

of Osten Sacken

(1881b, 1884)

and

independentlyarrivedattheconclusionthat

muscids

with

an

angular vein

M

and

bare

meron belonged

tothe

Anthomyaires/Anthomyidae and

notto the Tachinaires/Tachinidaeaspreviouslybelieved. In retrospect, ^itisapparent that Pandell6's proposals for the classification

of

the

Schizophora were

superior in several respects to those of Girscnner. Nevertheless, it

was

Girscnner's views, expressed succinctly

and accompanied by

clear

and

schematic drawings,that

had

the greaterinfluence

on

20th centurydipterol- ogy.

The

greateststimulustothe

work

ofthe early part

of

the20th century

came from

thebiological investigations

by

the

Russian

J.

A.

Portschinsky (1848- 1916). ^In

an

important pioneering paper publishedin

1885

(abstracted

and

discussedin

German by Osten

Sacken, 1887), Portschinsky

drew

attention to

an

apparent dual reproductivestrategy in

Musca

corvina[i.e.autumnalis]:

although normally laying

some 24

largeeggs, femalesⁱⁿthesouth of

Russia (Crimea and Caucasus)

^incubated

a

single large

egg and

^laid a 3rdinstar

maggot.

It is

now

clear, ofcourse, that

two

species

were

involvedhere, as Portschinsky himself subsequently recognized (Portschinsky, 1910).

He

continued withhisinvestigationsofoviparous

and

viviparous

Muscidae and

oftheircoprophagous, saprophagous,

and

carnivorouslarvae (Portschinsky, 1892, 1910, 1911),

and

hislater

work was reviewed by Townsend

(1911).

There

is

no doubt

that this

was enormously

influential in alerting field entomologists to the biological interest ofthisgroup, at

a

time

when

the

medical and

veterinary importance of Musca-flies

was

increasingly being recognized

and

this

gave a tremendous

impetustothe

fundamental work on

the

group

thatfollowed duringthe

1910s and

1920s.

Throughout

thisperiod(1885-1920)the

dominant

figureinthe

taxonomy

ofthe

Muscidae was

the

German

PaulStein (1852-1921),

who

specialized

almost

exclusively

on

this

group and who

single-handedly

brought

orderto the

chaos

into

which

the

group had sunk

duringthecourse ofthe19thcentury.

From

the

modern

standpoint, Stein'sconcept of genera

seems

rather limited, buthis grasp ofspecies

was

remarkable.

His

papers

were

methodical

and

comprehensive and

almost invariably contained keys;

he

travelled to the centersof

European

Dipterologyin

Germany,

England,

Sweden, and

France, tostudythetypes ofearlierauthors;and,towardsthe

end

ofhislife,

he was

ableto

produce a key

toallthe

European Muscidae

(Stein, 1916),

a key

to the

world

genera(Stein,1919),

and

catalogues ofthe

world

species(Stein,

Introduction 27

1919;in

Bezzi &

Stein, 1907). Stein

was most

interested inthe

"Anthomyi-

idae"(inthe19thcenturysense)

and

so

Musca and

alliedgenera

were

rather peripheraltothe

main

thrust

of

^his

work. He followed

Schiner (1862)in his concept

of

the

genus Musca. Reviewing

extensive collections

from

the Afrotropical

and

Orientalregions,

he was

thefirstto realize

how

large

and

diverse the

genus might prove

to

be

(and

how

very slipshod

and

inadequate

for.the

most

part hisprecursorsin tropical

Muscidae had

been).

For

thefirst time (Stein in

Bezzi &

^Stein, 1907) the Palaearctic

Musca

species

were

catalogued, with

an

inventory

of

recognized species,speciesincertae,

and synonyms, a

totalof

46 names.

The

first

two

decades

of

thecenturyalso

saw

theinitiationofseveralother strands

of work on Musca which

didnot

come

togetheruntil the

work

of Patton

and Malloch

inthe1920s.

The American

tachinid specialistC.

H.

T.

Townsend was always

interested inthe

importance of

reproductivebiology forsystematics (e.g.

Townsend,

₁₉₁₃₎

and on

severaloccasions

he

briefly turnedhis attention tothe

genus Musca. He was an

advocate

of

theconcept

of

restrictedgenera

and he

realized

from

Portschinsky's

account

of

Musca

corvina [autumnalis] thatthereproductivebiology

of

thisspecies

was

very different

from

that

of Musca

domestica.

He

erectedthe

new genus Eumusca

forthisspecies

(Townsend,

19_{1 1}).

A few

yearslater,afterstudying published

work on

theIndian

Musca

species,

he

erected the

genus Viviparomusca

for theviviparous speciesbezziiPatton

& Cragg (Townsend,

1915)

and

another

genus Awatia

forthe species indica

Awati

(=planiceps

Wiedemann) because

it

was a

larviparous

member of

the

Philaematomyia-group and had

well-de-

veloped

presuturaldorsocentralsetae.

Among

blood-sucking

Muscidae

collected in the first

decade of

the century

were

species

of Musca

that

could

abradevertebrateskin

and suck

blood,

but

atfirstit

was

notrealized that these

were

true

Musca

ratherthan biting

Stomoxyinae. Austen

(1909) described

a new genus Philaematomyia

(for

a

species subsequently

found

to

be

identical with crassirostris Stein)

which he

characterized as"a

remarkable

connecting-link

between

theordi-

nary

non-biting

Muscinae and Stomoxydinae." He noted

that the

powerful

proboscis

could

notpierce vertebrateskinbutthattheflies

used

the

enormous

prestomalteethtocut the skin

and

then

sucked up

blood.

He

did not consider

itto

be

particularlyrelated to

Musca.

Brunetti (1910) described

a new genus

Pristirhynchomyia (for

a

species subsequently

found

to

be

identical with

conducens Walker) which he

considered transitional

between Philaema- tomyia and Musca: he excluded both

genera

from

the

Stomoxyinae, though

without giving

any

reasons.

He

notedthathisspecies

was

also

a

"bitingfly,"

unabletopiercetheskinbut abletoenlargeexisting

wounds

orsores.

28 Systematic Database of Musca Names

Ithas

been

graduallyrealized thatthese featuresinbiology

and

structure areprogressive

and

thatvarious evolutionarystages arerepresentedwithin the

genus Musca,

astheyareinother genera

of

theMuscini.

The

fragmenta- tionofthe

genus on

thebasis

of a few autapomorphies

resultsin

a

seriesof typological units ratherthan

monophyletic

groups

and does

notreflectthe evolutionaryhistory

of

thegenus.

Work

inIndia

by Awati (1915-1920) and

Patton

& Cragg

(1912, 1913a, b)increasedthe

number

ofspecies

and paved

the

way

for

advances made

in the1920s. Awati's

work was

particularlyimportant, since

he looked

in

such

detailat

a wide

range of morphological, anatomical,

and

biological features, but the lack of

any

reference material

from

his

work

has

hampered

the recognition ofthe actual species that

he

discussed: indeed, it is stillnot possibleto interpretallthe

names

that

he

proposed. Patton did

an enormous amount

of fieldwork

and

breedingof

Musca

speciesin

South

India

and

this borefruit

when he

returnedto

England and began

extensivestudies

on

the

taxonomy and

systematics

of Musca: he was

ableto

combine

hisfirst-hand

knowledge of

reproductivestrategies,biology,adultfeedinghabits,etc,with his systematic researches.

Both

authors

used Musca

in the

broad

sense defined

by

Stein.

The Modern Era (1921-1992)

A

^conflict

^between

^generic"lumpers"

and

"splitters,"familiar in

most

groups

of

insects,

developed

duringthe1920s,but

a

carefulstudy of various character-suites

among

thespecies

of Musca and

abetterunderstanding of theevolutionarytrendswithinthe

genus soon gave

riseto

a

delimitationof the

genus which

has

remained

stable

now

^for

60

years.

The

varioussplinter genera defined duringthe

1920s

arerecognizedatthe levelof

subgenera

or speciesgroups.

The

period

up

to

1937 was dominated by

the Briton

W.

S. Patton (1867-1960).

Through

hisfirstcareer in theIndian

Medical

Service,Patton

developed an

interest in all the higher flies

of medical and

veterinary importance(especially

Muscidae and

Calliphoridae)

and

these earlyyears of practical

work underpinned

all his subsequent systematic research. His correspondence

and exchanges

with other

muscoid workers

ofthe

day

(E.E.

Austen,J.Villeneuve, J.

M.

Aldrich,

and

especially

M.

Bezzi)

showed him

thatthe

same

species

were

often

masquerading under

several different

names

and

thatthe

names

themselves

were

oftenappliedin

a

quite

haphazard and

inconsistent

manner.

Believing that

he

understoodthe species

and

species limits sufficiently well,

he

studiedthetypes

and

othermaterialin

European

museums and

private collections (Patton,1922a, 1923a, 1923b, 1925).

From

Introduction 29

thisbasis

he proceeded

topublishrevisions

of

theOrientalspecies(Patton

&

Senior-White, 1924;Patton, 1937a,b), Afrotropical species (Patton 1926, 1936),

and

Palaearctic species (Patton, 1933b,c).

He

also

undertook more fundamental comparative

studiesofthe

male

genitalia (Patton,1932, 1933a), theantennae (Patton

&

MacGill, 1925), theevolutionofthe

bloodsucking

habit (Patton, 1934),

and

supervised

a comparative

study

of

the

female

ovipositor(Ho, 1938).

Patton

was convinced

that

Musca

as

he understood

it(and asStein

had understood

it,

and

as the present generation still understands it)

was an

indivisible

monophyletic

unit

He

put

forward

characters to support this contention,

such

as the

autapomorphous

presence

of a

large

sensorium

atthe

base of

the antennal basoflagellomere (Patton, 1932)

and

also

adduced arguments

to

show

thatthe

more

strikingbiological features in

Musca

(such asviviparity

and

modificationsinthe

female

reproductive system, adapta- tions inthe

male

genitalia,biting

and

bloodsucking)areprogressive and,in thecase

of

the blood-feeding,

have

arisen

more

than

once

withinthegenus.

He

recognized only three

groups

in the genus, the domestica-group, the sorbens-group,

and

the lusoria-group. In discussion

he was

ableto

persuade Bezzi

that this

was

the right course,

though Bezzi had

earlierjoinedin the fragmentation process

begun by Townsend. He

dismissed

Townsend's

splin- ter

genera

withderision.

But he could notreach any

accord with

Malloch and

the

two remained

irreconcilable.Itisclear

now

that

even

Patton misidentified

a

greatdeal

of

^material:throughouthislong career

he

re-identified,mis-iden- tified,

and changed

his

mind about

specieslimits

and names and

his

keys

are generallypoor,

based on

rather

weak

characters

and

difficultto use.

Malloch

too misidentifiedspecies.Itis

obvious

that

much of

^their

argument over

detail

can be

resolved

once

it is

known what

species they actually

meant by

particular

names:

for instance,

both had iogmatic views over where

"planiceps

Wiedemann"

should

be

placed,yetneither correctly identifiedthis species and,asthey

were

probably not

even

discussingthe

same

species,their discord appears completelyfutile.

On a

higher systematiclevel,the

argument

resolves itself

over

the ranking

of

hierarchies,

whether

as species groups (Patton) or

genera

(Malloch).

The

present system, largely

worked

out

by Emden

(1939, 1965), represents

a

synthesis

of

the

work

of Patton

and

Malloch.

The

Italian

Mario Bezzi (1868-1927) worked on Musca

atvariousstages

of

his career,

mainly

describing

new

species.

He gave a key

to all the Mediterranean

and

Africanspecies,

and

recognized

a number

of

subgroups

(Bezzi, 1911).

Turning

to theblood-sucking

forms

(Bezzi,1921)

he showed

that the species

of Musca

sensu Stein

were now

distributed

among two

30 Systematic Database of Musca Names

subfamilies,

Philaematomyiinae {Philaematomyia,

Pristirhyncomyia, Ptilolepis)

and Muscinae

(5 genera).

He

subsequently (Bezzi, 1923) de- scribed another subgenus,Lissosterna, for the desert species

Musca

albina

Wiedemann.

The work of

the Scottish-American

John R. Malloch (1875-1963)

ad-

vanced knowledge

ofthefamily

Muscidae

to

a

quite

remarkable

degree

and

in

two

studiesofthe

genus Musca

(1924, 1925;furthernotesin1929b, 1932)

he showed

theinnovative

approach and

remarkableabilitytoperceive

new

characters that are evidentinallhis

work. He

discoveredseveralvaluable features for identifying

and

classifying speciesof

Musca, some

of

which were

adopted,albeit reluctantly,

by

Patton.

He

too

was

uncertain

how

to

rank

the

groups

that

he

coulddefine: "Itisdifficulttoassign

a

definite status to the segregates hereinaccepted,^for,while

some workers would

consider

them

as valid genera, others

would

just as certainlyrank

them

at

most

as subgenera, or

even

refuse toaccept

them

as

named

divisions"(Malloch, 1924:132).

He

proceeded

tolist

them

asgenera(1924),thenassubgenera (1925) but with

Philaematomyia and

Ptilolepis as separate genera,

and

thenfinallyasfull genera again (1929b), recognising

a

total of 8 genera {Musca,

Byomya,

Lissosterna,

Plaxemya, Philaematomyia, Eumusca, Viviparomusca,

Ptilolepis).This

work was

rejected

by

Patton,

who

regardeditas

a

falsifica- tionofthe true relationshipswithin

Musca.

Since

both were

handlingin part misidentified material, it

was

not possible for

them

tofind

any common

ground

at all,

though

subsequent

work

has

acknowledged

merit in

both

approaches.

Two

contrasting

views

of

Musca appeared

ⁱⁿ 1937.

Townsend summa-

rized his

views on

thegenera of the

Muscini and

divided

Musca

intoeleven genera:Awatia,

Byomya, Eumusca,

Lissosterna,

Musca, Philaematomyia, Plaxemya,

Pristirhynchomyia, Ptilolepis,

Sphora, and Viviparomusca (Townsend,

1937).Sincethesesegregates

were based on

thestudy

of a

very

few

species

and were founded

often

on

biologicaloranatomicalfeatures,they

have

not

been

accepted

by

anyone.In the

same

year,

Seguy

(1937)

produced a volume

^for

Genera Insectorum

that

gave a

catalogue

of World Muscidae.

He

followed Malloch's(1925) concept

of Musca,

but

ranked

theblood-suck- ingPtilolepis

and Philaematomyia

asdistinctgenera.

For

these threegenera

he

catalogued

a

totalof

92

^species,with

164 names

(valid

and synonymous).

The 92

species include

a number

thatare

no

longer recognizedin

Musca,

species incertae, misidentifications,

and names

that are

now synonyms,

whilstseveral

names

that

do

in fact

belong

to

Musca were

overlooked

by

S6guy.

Introduction 31