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Contributions

of the

American Entomological Institute

Volume 3, Number 1, 1968

MOSQUITO STUDIES (Diptera, Culicidae)

VII. The Culicidae of New Zealand.

By John N. Belkin.

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Contributions

of the

American Entomological Institute

Volume 3, Number 1, 1968

MOSQUITO STUDIES (Diptera, Culicidae)

VII. The Culicidae of New Zealand.

By John N. Belkin.

MOSQUITO STUDIES (Diptera, Culicidae)

VII. THE CULICIDAE OF NEW ZEALAND!

John N. Belkin2

CONTENTS

INTRODUCTION

LAMY oo ea ee a a

SYSTEMATICS AND BIOGEOGRAPHY.

KEYS TO SUBFAMILIES.

SUBFAMILY DIXINAE.

Keys to Tribes and Genera.

Tribe Dixini . ‘

Genus Neodixa .

1. Neodixa minuta.

Genus Nothodixa

Keys to Species

2. Nothodixa campbelli

3. Nothodixa philpotti .

4. Nothodixa otagensis. ‘

0. Nothodixa septentrionalis .

Tribe Paradixini .

Genus Paradixa.

Keys to Species

. Paradixa neozelandica

. Paradixa fuscinervis .

. Paradixa tonnoiri, n. sp.

. Paradixa harrisi .

Oo CO-~I1

I this investigation was supported by National Science Foundation Research

Grants G18961 and GB2270.

Oh oa piiient of Zoology, University of California, Los Angeles, Califor-

nia 90024.

2 Contrib. Amer. Ent. Inst., vol.

SUBFAMILY CULICINAE

Keys to Trib

es and Genera.

Tribe Culicini

Genus C

ulex, subgenus Culex Culex

Tribe Culise

Genus Culiseta, subgenus Climacura :

Culiseta (Climacura) tonnoiri.

14.

Tribe Manso

Genus C

15. Coquillettidia (A. ) tenuipalpis.

16.

Keys to Species. . .

. Culex (C.) pervigilans .

. Culex (C.) rotoruae, n. sp.

. Culex (C.) asteliae, n. sp..

. Culex (C.) quinquefasciatus.

tini

niini.

oauilietid =.

Keys to Species .

$, nO. 1, 1968

Subgenus Austromansonia, oe ‘subg.

Subgenus Coquillettidia.

Coquillettidia (C.) iracunda.

Tribe Aedini . : :

Keys to Genera, Subgenera and ‘Species ;

Genus Opifex.

17.

Genus A

18.

19.

20.

21.

22.

Opifex fuscus .

Aedes . : :

~ Subgenus Nothoskusea :

Aedes (N.) chathamicus

Subgenus Halaedes.

Aedes (H. ) australis.

Subgenus Ochlerotatus .

Aedes (O.) antipodeus .

Aedes (O.) subalbirostris

Subgenus Finlaya .

Aedes (F. ) notoseriptus ;

Tribe Sabethini .

Genus Maorigoeldia .

23.

Maorigoeldia areyropus

SUBFAMILY CHAOBORINAE.

Tribe Corethrellini .

Genus C

EIGURES: 2c.

APPENDIX A

orethrella

Subgenus Notocorethrella, n. Ae

24. Corethrella (N. ) novaezealandiae ‘

RECORDS OF EXTRALIMITAL SPECIES

REFERENCES CITED.

NZ Collection Data .

APPENDIX B

Laboratory Colonization of Paradixa neozelandica Tonnoir

By Don

INDEX: oc.

Forsyth

180

Belkin: Culicidae of New Zealand 3

INTRODUCTION

In a previous study (Belkin 1962:15-18) I alluded to the extraordinary na-

ture and taxonomic importance of the mosquito fauna of New Zealand and out-

lying islands. All but 1 of the indigenous species of this fauna are endemic and

the majority of these appear to have retained more significant primitive fea-

tures than any other living representatives of their respective phylads. The

desirability of a thorough knowledge of this relict mosquito fauna from the

standpoint of composition, distribution and detailed knowledge of the individual

Species cannot be overemphasized for studies on the morphology and phylogeny

of the Culicidae and their biogeography. Unfortunately even at the conclusion

of the above-mentioned study only very fragmentary knowledge of this fauna was

available and very little material for more detailed studies was to be found in

museums outside of New Zealand.

The present study is based primarily on material collected by myself and

Donald A. Schroeder in 1963-1964. Additional specimens have been kindly pro-

vided by L.J. Dumbleton, D.A. Forsyth and E.R. Nye. I have also examined

the collections at the Auckland Museum (AUCK), Canterbury Museum (CANT)

and at the Department of Scientific and Industrial Research, Nelson (NELS). I

thank all the above individuals and institutions and also Roy Elliott, E.S. Gour-

lay, D.D. McCarthy, J.S. Pillai and particularly C. Arthur and Mary K.

Schroeder for innumerable favors and assistance during my brief visit in New

Zealand. I am also indebted to Patricia Donnelly for a preliminary study of the

larval and pupal chaetotaxy of Nothodixa campbelli and Paradixa fuscinervis,

carried on as a research problem in the Department of Zoology, University of

California, Los Angeles. Thomas J. Zavortink, O. George W. Berlinand Sandra

J. Heinemann prepared and checked most of the material for this study. The

preliminary drawings were made by Rainer Beck and Anne Acevedo and the

final drawings were all made by Sharon Burmeister. Without their arduous toil

this study could not have been completed and I am most grateful to all of them

for expert assistance.

Nearly 20, 000 specimens (1376 o, 1844 9, 13662 larvae, 2706 pupae), rep-

resenting all the 24 recognized species and including more than 1000 individual

rearings (580 larval, 364 pupal, 100 incomplete) of 19 species, were examined

during this study. Some material of both sexes and all stages was studied for

all species except as follows: (1) no females or immature stages of Neodixa

minuta, (2) no males or immature stages of Nothodixa philpotti, (3) no imma-

ture stages of Nothodixa otagensis, (4) no females of Nothodixa septentrionalis,

(5) no larvae of Coquillettidia tenuipalpis, (6) no larvae of Coquillettidia ira-

cunda, and (7) no immature stages of Corethrella novaezealandiae.

The primary objectives of this publication are to describe and figure in

some detail previously unrecognized species or stages and to provide more

complete descriptions and illustrations of those New Zealand species which are

the types of generic group taxa. Special emphasis is given to the morphology

of the poorly known Dixinae. I have also taken this opportunity to present a re-

view of previously published data on New Zealand Culicidae to provide a basis

for more detailed future studies which I hope will be undertaken by New Zealand

investigators.

The methods of study and presentation are essentially similar to those I

used in ''The Mosquitoes of the South Pacific'’ (Belkin 1962). A few modifica-

tions in the terminology are introduced and explained in the general discussions

of the Dixinae and Culicinae. In the illustrations only some pertinent details

4 Contrib: Amer. .Ent. Inst.,. volv 3, no. 1, 1968

are shown and many features have been eliminated. This applies particularly

to illustrations of adults where the scale patternis usually not shown and to the

figures of leg segments which are designed merely to indicate the proportions.

Because of paucity of published information of the bionomics and distribu-

tion of New Zealand Culicidae I have included an Appendix in which the data on

our collections are summarized and have provided detailed distribution lists

and maps for each species. The collection data are not repeated in the lists

but can be obtained from Appendix A by reference to the collection number

which is identified by the code NZ. The numbers on the detailed distribution

maps also refer to the NZ collection numbers in Appendix A.

In the distribution lists, the arrangements of localities is from north to

south and west to east, grouped under the land districts except for the islands

of Hauraki Gulf which are listed individually after the North Auckland land dis-

trict. Included in these lists are all other records of material I have examined

as well as some published records of specimens I have not seen but consider

reliable. All these miscellaneous records are indicated on the detailed distri-

bution maps but without collection numbers.

MORPHOLOGY

Basic to phylogenetic studiesis the determination of primitive (plesiomorph)

and derived (apomorph) states of various attributes. In this respect, the culi-

cids of New Zealand hold a unique position because in nearly every indigenous

phylad many primitive characters have been retained and because both the Dix-

inae and Culicinae are well represented. As one of the main criteria available

for determining the primitive states in a group without a good paleontological

record is comparison with the conditions exhibited in coordinate taxa, I have

made a special effort to study the morphology of the Dixinae in order to deter-

mine the probable primitive states of various attributes in the Culicinae.

Due to the lack of material from Southeast Australia, Tasmania, Chile and

Patagonia, thorough detailed comparative morphological studies could not be

carried out during this investigation. However, the preliminary studies on the

Dixinae of New Zealand (see) suggest that this group holds the key to under-

standing the morphological composition of the male genitalia of the Culicinae as

has been shown to be the case by Martini (1923) for the morphology and chaeto-

taxy of the caudal abdominal segments of the larvae. I have proposed a new

morphological terminology for the components of the phallosome of the Dixinae

which is discussed under that subfamily and is illustratedin fig. 7. One of these

components, the opisthophallus, is recognized here for the first time in the Cu-

licinae (See). Detailed comparative studies of the immature stages of the Dix-

inae of New Zealand have led to some corrections and a reinterpretation of the

chaetotaxy with the result that the pattern is so similar to that of the Culicinae

that there is little doubt that it is truly homologous in the 2 groups.

In the discussion of the Culicinae (see), I have indicated what I believe to

be primitive states for some characters of this group exhibited by New Zealand

species. Almost all of these are similar to those of New Zealand Dixinae but

some are unique developments of the Culicinae.

Belkin: Culicidae of New Zealand s)

SYSTEMATICS AND BIOGEOGRAPHY

Although our current knowledge of the culicid fauna of New Zealand is still

fragmentary some speculations regarding its affinity, originand evolution seem

warranted. I have summarized my views on these matters in the discussions

of the Dixinae and Culicinae, which are both well represented in New Zealand.

The details of the probable affinities are dealt with under the individual species

and groups. Here only a general summary and broad generalizations are pre-

sented.

Of the 24 species definitely known to occur in New Zealand, Culex (C.)

quinquefasciatus is undoubtedly a recent introduction and Aedes (F.) notoscrip-

tus was probably alsointroduced subsequent to the colonization of New Zealand.

Of the remaining 22 species, only Aedes (H.) australis is known outside of the

New Zealand area. Three monotypic genera (Neodixa, Opifex and Maorigoel-

dia) and 3 monotypic subgenera (Austromansonia, Nothoskusea and Notocoreth-

rella) are recognized for New Zealand species. Only a few of the other endemic

Species appear to have close relatives anywhere in the world.

The culicid fauna of New Zealand is a well balanced one, with a represen-

tation of all 3 subfamilies and all the major phyletic lines within each of these

except the Chaoborinae. The poor representation of the latter may be due at

least in part to the predaceous nature of the larvae. Nearly all the major phy-

letic lines present in New Zealand appear to be ancient ones and within these,

the individual elements seem to have retained a greater number of signifi-

cant primitive features than any other living representatives of their respective

phylads. The apparent absence of certain groups generally considered to be

"primitive, '' especially the tribes Anophelini, Toxorhynchitini and Uranotaeni-

ini, is very striking. I submit that these phylads failed to invade New Zealand

either because of their absence in the source areas or their representation

there by species with poor dispersal powers or limited adaptability to new en-

vironments at the time that dispersal routes were available. Neither the Tox-

orhynchitini nor the Uranotaeniini are now known from Southeast Australia-

Tasmania. The Anophelini are represented by a number of apparently ancient

relict Species in these areas and it is possible that they did reach New Zealand

but have not been able to survive.

A striking feature of the fauna is the presence of only 2 indigenous contain-

er breeders, Maorigoeldia argyropus and Culex (C.) asteliae, all the remain-

ing indigenous forms being ground water breeders. It appears that the spe-

cialized container habitat was first utilized in the Culicinae by the Sabethini

which show the most striking derived features in the larval stage and are all

container breeders. The utilization of this habitat appears to beincipient in the

ancient stocks of Culicini as shown by the facultative use of large treeholes by

many species of Culex (Culex) and the derivation of Culex (C.) asteliae from the

pervigilans stock in New Zealand. The absence of indigenous container breed-

ers in the Aedini of New Zealand suggests that this habit had not been acquired

by the more ancient stocks of the tribe at the time of its entry of New Zealand.

The majority of New Zealand species appear to have moderate powers of

dispersal and great ability to survive in small populations as shown by their

presence on small islands in Hauraki Gulf, such as Little Barrier Island.

There is relatively little geographical differentiation in the culicid fauna of

New Zealand; most species occur throughout the main islands and adjacent is-

lets. The significant exceptions are: (1) Aedes (N.) chathamicus, restricted to

Chatham Island, (2) Aedes (H.) australis and Aedes (O.) subalbirostris, re-

6 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

stricted to the southern part of South Island, and (3) several relict species in

the Dixinae (see). In the widespread species, there is little indication of differ-

entiation in the different populations except to a limited degree in Culex (C.)

pervigilans.

In general, each phyletic line is represented by a single species in New

Zealand. The only exceptions are in the Dixinae(see) and the Culex pervigilans

complex. Speciation within New Zealand in both these instances appears to

have occurred through isolation, accompanied by ecological (Culex, Nothodixa).

or genitalic (Paradixa) barriers.

The affinities of the culicid fauna of New Zealand are largely with the fauna

of Southeast Australia-Tasmania. Only the genera Nothodixa and Neodixa show

definite relationship with the fauna of both South Chile-Patagonia and Southeast

Australia-Tasmania. The affinities in all but 1 or 2 cases are not very close

and in nearly all these instances the New Zealand species have retained more

primitive features than their relatives and therefore it appears that they could

not have derived directly from the latter. In Several instances, apparent rela-

tives of New Zealand stocks, and in some cases Australian as well, are found

in the Northern Hemisphere, notably in Paradixa, Culex (C.) pervigilans, Cu-

liseta (Climacura), Coquillettidia (C.) and Aedes (Ochlerotatus).

At least 3 waves of dispersal probably took place in the population of New

Zealand by culicid stocks. The earliest involved only the Nothodixa-Neodixa

line and occurred probably at the same time as the classical southern hemi-

sphere dispersals of plants. Evidently the other culicid stocks were not pres-

ent in the source areas at this time. The second dispersal involved the bulk of

the culicid fauna of New Zealand after the connection (direct or indirect) with

South America was broken. There is a suggestion that 2 separate waves may

have been involved in this dispersal in the separation of Opifex and Aedes (N. )

chathamicus, the presence of different lineages in Paradixa, and the slight dif-

ferentiation between the elements on North Island and South Island. The final

dispersal is a much more recent one, involving apparently only Aedes (H.) aus-

tralis.

In summary, the indigenous culicid fauna of New Zealand has all the ear-

marks of an ancient continental fauna, probably largely Mesozoic in origin.

The land connections for the dispersals need not but may have been continuous;

relatively short barriers may have been efficient filters at certain times and

may account for the absence of a few ancient stocks. On the other hand, the

barriers probably were not very great at other times as the Sabethini and the

Dixinae, which are apparently incapable of crossing extensive ocean barriers,

were able to reach New Zealand. Subsequent to the isolation of the New Zea-

land area and coincident with later geological changes a limited geographical

differentiation of the original Mesozoic culicid fauna has taken place without any

additions (except perhaps for Aedes (H.) australis) from external sources until

the introduction through human agency of Culex (C.) quinquefasciatus and prob-

ably Aedes (F.) notoscriptus. In the course of time there has probably been

considerable attrition in the culicid fauna, only those species surviving that

have been able to adapt to the great environmental changes since the Mesozoic.

This, I believe is shown by the great plasticity of the majority of present day

forms in their ability to utilize a wide range of habitats and in their considera-

ble tolerance of temperature and salinity.

As I have indicated earlier (Belkin 1962:58), the most likely places for the

ultimate origin of the stocks which populated New Zealand and the other South

Pacific islands are islands in the tropical unstable Old World intercontinental

area.

Belkin: Culicidae of New Zealand 7

Even the so-called antarctic element represented by Nothodixa-Neodixa

could have had its ultimate origin in this area and dispersed only to the south

because of the availability of dispersal routes only in this direction. This in-

tercontinental origin of new types would explain also the presence of related but

distinct stocks in the northern and southern hemispheres, as in the case of Pa-

radixa, Culex (C.) pervigilans, Culiseta (Climacura), Coquillettidia (C.) and

Aedes (Ochlerotatus), by the dispersal of a new type from the intercontinental

area to both hemispheres because of availability of dispersal routes and subse-

quent independent evolution in widely separated areas.

2(1).

K# YS TO SUBFAMILIES

Adults

Mouthparts long, labium forming a long rigid proboscis. . . Culicinae

Mouthparts short, labium not elongated into a rigid proboscis ... . 2

Wing veins without or with only a few inconspicuous short hairs; vein Sc

ending on wing margin near level of furcationof R.. .. . Dixinae

Wing veins with numerous long hairs or hairlike scales; vein Sc ending

on wing margin distinctly distad of furcation of R (24. Corethrella no-

VOGVERISNOIGE). obs bye ai es 2k ec sw we bo ee ee

Male Genitalia

A distinct paramere developed between the basal piece and aedeagus

ever a ge Seog ar ae gs Se ce Culicinae

Aedeagal sclerites articulating directly with basal piece. ...... 2

Phallosome very simple, consisting of a poorly developed basal piece

and projecting aedeagal sclerites, which are not bent back into an

aedeagal pouch (24. Corethrella novaezealandiae). . . . Chaoborinae

Phallosome complex, basal piece strongly developed, prosophallus or

opisthophallus or both strongly developed, aedeagal pouch always pres-

ent, penis filament sometimes developed ....... . . Dixinae

Pupae

Abdominal tergites with incomplete submedian transverse ridges; seg-

ment IX well developed, distinct sternally. .... .. .. Dixinae

Abdominal tergites without submedian transverse ridges; segment IX

poorly developed, mdistinct sternally ... svc os en eed evn eee

Paddles fused at base, not articulated or trumpet without pinna or trach-

eoid but with large ovoid reticulate meatus and small apical process

ee Chaoborinae

Paddles always separate and AMieniaeen at base and trumpet usually with

pinna; if pinna indistinct then trumpet without large ovoid reticulate

meatus and small apical proceSS:.: augue wisn 25 5 oe ULC mae

Larvae

Venter of abdominal segments I or I and II with short unsegmented paired

8 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

lobes (prolegs) ending in recurved spines; thoracic segments same

width as abdominal and rather distinctly separated. ... . Dixinae

Venter of abdominal segments I and II without such lobes; thoracic seg-

ments consolidated into a single mass wider than abdominal segments

ANG MOL -GOCMENLCG tis sic e: cur alte Merrie oe ine aaa tel be os badly 6 Laie me

2(1). Mouthbrushes completely absent or aol cache at most by about 5 long

bristlelike spicules... . Kowa > oe CMpObOrinae

Mouthbrushes always well jeveloned. asually os numerous, rarely re-

duced to about 10 heavy long flattened spines. . ... . .. .Culicinae

SUBFAMILY DIXINAE

The Dixa Midges

Dixa midges are readily recognized from true mosquitoes by the charac-

ters given in the key to the subfamilies. A general account of the subfamily

with emphasis on New Zealand forms is given by Belkin (1962:93-96) to which

the following additions and corrections based on a more detailed examination

during the present study should be added.

MALE GENITALIA. A thorough comparative study of the male genitalia of

the Dixinae has not been made yet but it appears from even a cursory examina-

tion of the New Zealand species and a few extralimital forms that the Dixinae

hold the key tounderstanding the morphological composition of the male genitalia

of the Culicinae. There is a bewildering array of specializations of the basic

sclerotizations and apodemes associated with the genital opening, intromittent

organ, Sidepiece and proctiger developed from a very generalized primitive

type which is perhaps best represented in the New Zealand species of Nothodixa

and Paradixa.

For the present I am continuing the terminology I have used in the past

(Belkin 1962) and I am introducing only a few new terms which are discussed

here and illustrated in fig.7. In this discussion the usual descriptive taxo-

nomic terminology of orientation will be used and the morphological relation-

ships will be mentioned only when they are obvious or necessary.

The basic support and articulation points for all sclerites associated with

the phallosome complex and proctiger are provided directly or indirectly by a

highly complex apodeme, the basal piece, which appears to be primarily a de-

velopment from the mesal dorsal angle of the sidepiece, although primitively

it may have been an apodeme from the entire mesal margin of the sidepiece or

from both its dorsal and ventral mesal angles. In the Dixinae the basal piece

is much more extensive and complex than in most Culicinae. I distinguish 2

parts in the basal piece, the distal division (ventral) which projects inside the

sidepiece itself and the proximal division (dorsal) which projects inside seg-

ment IX. The 2 divisions are continuous and it is not always possible to sepa-

rate them distinctly; certain portions of either division may acquire articular

points.

Although very complex, the phallosome of New Zealand Paradixa seems to

be the most primitive in the entire family. I recognize 3 basic components:

(1) a ventral prosophallus or claspette, (2) the phallus proper and (3) a dorsal

opisthophallus or dorsal claspette.

The prosophallus consists of a transverse lobe between the mesal ventral

edges of the 2 sidepieces. It is supported by a simple transverse sclerite

Belkin: Culicidae of New Zealand 9

(prosophallic sclerite) which is joined on each side to a ventral arm of the dis-

tal division of the basal piece. The prosophallus is probably the homolog of

the claspette of the Culicinae. In the Dixinae it is variously modified into me-

dian or submedian processes along its distal margin and may be separated into

basal and distal portions which, however, are continuous. Ventrad of the pro-

sophallus is a small depression (sternal pocket) between the posterior margin

of sternite IX and the base of the 2 sidepieces which meet at the bottom of the

depression.

The phallus is the median lobe associated with the gonopore. In New Zea-

land Paradixa its support is provided by a sclerite (aedeagal sclerite) on each

side connected to the central part of the proximal division of the basal piece.

In other Dixinae (e.g. N. Z. Nothodixa) it becomes articulated with a long pro-

cess from the latter. This process through further differentiation and articu-

lation, I believe, becomes the paramere of the Culicinae. The aedeagal scle-

rites proper form the aedeagus and may become completely fused on the dorsal

surface butare usually at least partially separated on the ventral surface of the

phallus. :

The gonopore in New Zealand Paradixa is always ventrad (morphologically

cephalad) of the aedeagal sclerites. It is usually in a depressed membrane be-

tween the aedeagal sclerites and the prosophallic sclerite but in harrisi it is

located within the apex of the proximal division of the prosophallic sclerite. It

is possible that the latter condition is the primitive one but this cannot be re-

solved without extensive comparative study of related families. I have noted in

some North American Dixinae similar sclerotizations around the gonopore ap-

parently derived from a proximal division of the prosophallus.

The opisthophallus consists of a transverse lobe between the mesal dorsal

edges of the 2 sidepieces. It is dorsad (morphologically caudad) of the phal-

lus and lies between the latter and the venter of the proctiger. I believe that

the opisthophallus belongs to the 9th abdominal segment but have made no at-

tempt to determine its exact morphological composition. The opisthophallus

is supported on each side by a sclerite (opisthophallalic or lateral sclerite)

articulated with the outer part of the proximal division of the basal piece. In

New Zealand Paradixa the opisthophallus is very complex and its lateral scle-

rite articulates dorsally with the ventrolateral sclerite of the proctiger. In

New Zealand Nothodixa the opisthophallus is a relatively simple large lobe

whose lateral sclerites are poorly sclerotized and do not articulate with the

membranous proctiger distally. The opisthophallus has not been studied or

even recognized in the majority of Culicinae. However, it is evident that it is

strongly developed in some forms, notably in Maorigoeldia and in the subgenus

Verrallina of Aedes. :

The phallic complex in New Zealand species of Nothodixais of considerable

interest Since it appears to represent a very primitive condition which may

explain the derivation of the penis sheath and penis filament in other species of

Nothodixa and various extralimital paradixines. In the New Zealand species

the parameres are very long slender capitate apodemes which project caudo-

mesad to articulate with a moderately sclerotized cordate median plate located

at the dorsal end of a median longitudinal depression bounded at the ventral end

by the prosophallus and supported laterally by a pair of submedian longitudinal

lightly sclerotized processes from the prosophallus. This depression is prob-

ably homologous with the aedeagal pouch of Spielman (1964, 1966) and the

phallotheca of Snodgrass (1935). The cordate median plate I consider to be

composed of fused aedeagal sclerites. At the apex (anterior end) of the cor-

10 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

date plate is a small scoop-shaped, rounded, heavily-sclerotized median pro-

cess which is continued ventrally into the aedeagal pouch as a short slender

curved free median penis filament. This filament appears to have an open

groove on one surface and it is probably the actual intromittent organ which

probably enters the distal end of the spermathecal duct of the female in copu-

lation. I think it is very likely that the penis filament is also developed from

the aedeagal sclerites. It is possible, however, that the apical process of the

cordate plate is derived from the proximal division of the prosophallus, that it

surrounds the gonopore as in P. harrisi, and that it is joined to the aedeagal

sclerites secondarily. I have not been able to determine exactly the relation-

ship of the gonopore to the penis filament but it is probably located in the vicin-

ity of the apical process of the cordate plate which bears a striking resemblance

to a structure found at the ventral base of the aedeagus near the gonopore in

many Culicinae.

Nothodixa chilensis Alexander, 1913 shows a further step in the evolution

of the penis filament and the development of the penis sheath. In this species

the parameres are very Short, transverse and heavy and are broadly joined to

the heavily-sclerotized posterolateral walls (aedeagal sclerites) of the aedea-

gal pouch which is very deep and narrow. The base (anterior end) of the

aedeagal pouch is developed into a very narrow invagination (the penis sheath)

projecting into the body. The penis filament arises at the fused bases of the 2

aedeagal sclerites and projects into the invagination cephalad along its dorsal

wall then curves back distad along its ventral wall and finally dorsad between

the narrowed lips of the aedeagal cup to emerge free a considerable distance

above its base. The proximal part of the penis filament appears to be solid

and part of the posterior wall of the aedeagal pouch. Therefore it is possible

that the true penis filament is the free distal, grooved part only.

Two South American species of Nothodixa (ensifera and nitida, both Ed-

wards, 1930) are noted for the development of an external penis sheath con-

taining the penis filament; in ensifera this sheath is longer than the entire ab-

domen. I have not studied the morphology of the phallosome of these species

but it seems probable that the penis sheath of these species is nothing more

than an external evagination of the distal part of the aedeagal pouch which in-

stead of projecting into the body, has been evaginated in the area dorsad of the

prosophallus to accommodate the excessively long penis filament.

Elongate penis filaments located in internal sheaths in general similar to

but often much longer than in Nothodixa chilensis are known in a number of

paradixines, e.g. californica Johannsen, 1923 and Dixina solomonis Belkin,

1962. In californica the dorsal and ventral walls of the penis sheath are

strongly sclerotized, these sclerotizations being continuous with those of the

base of the aedeagal sclerites and of the submedian processes of the prosophal-

lus. The free penis filament arises from the ventral wall deep within the penis

sheath and curves along the ventral wall. Peters and Cook (1966) and others

have used incorrectly the term "ejaculatory duct" for the penis filament of the

Dixinae. 3

It is of interest that the intromittent organ (aedeagus, penis filament) in

the Culicidae is withdrawn at rest into a pouch (aedeagal pouch, penis sheath)

and that the gonopore lies nearits base on the ventral surface (morphologically

anterior) and not at its apparent apex. I suggested (Belkin 1962:178) that in Cu-

lex the aedeagus proper is deeply retracted at rest and Spielman (1964, 1966)

has demonstrated this and shown the relationship of the gonopore to the aedea-

gus in Aedes aegypti and Culex quinquefasciatus. It appears that in most Culi-

Belkin: Culicidae of New Zealand 11

cinae the aedeagus is not a tubular structure but is troughlike on its ventral

surface which is largely membranous and that within this membrane lies the

gonopore. I have shown above that in New Zealand Nothodixa the penis filament

is withdrawn at rest deep into the aedeagal pouch. In New Zealand Paradixa the

aedeagal sclerites are also withdrawn at rest into a shallow depression between

the sidepieces and the prosophallus and are folded back against a median pro-

cess of the distal division of the prosophallus so that the gonopore lies hidden

in a small pocket ventrad of them.

The proctiger of the Dixinae also shows much differentiation which may be

of taxonomic significance. In many forms (e.g. New Zealand Nothodixa, many

other Dixini and apparently some Paradixini) it has no distinct sclerotizations

and the cercal lobe is large and bears many setae. In New Zealand Paradixa

strong basolateral sclerotizations with spinelike processes are developed and

are articulated at base with the lateral sclerite of the opisthophallus. In this

group the cercal setae are few in number and are borne on a distinct process

which may be short or very long.

The ninth segment is strongly developed in all Dixinae and exhibits much

differentiation in various groups. Its tergite is greatly enlarged in species or

groups in which the opisthophallus and the proctiger are strongly developed,

as in New Zealand Paradixa. In the latter, the lateral tergite of the opistho-

phallus appears as a posterior lateral piece of abdominal tergite IX.

The sidepiece and clasper are the only parts of the male genitalia of the

Dixinae which have attracted wide attention in the past. Because of very strong

Specific differentiation in the size, shape and lobes of these structures, the

Sidepiece and clasper have been the primary means of recognizing and diag-

nosing species and most workers have not gone beyond this point in studying

the genitalia. None of these specializations appear to have any significant

value in recognizing natural groups or evolutionary lines.

FEMALE GENITALIA. The female genitalia of the Dixinae of New Zea-

land have not been studied in detail and little attention has been paid to them

elsewhere until the work of Peters and Cook (1966). These authors figured

them for many North American species but did not attempt to use them in the

classification of the group. There appear to be many different developments

in the bursa copulatrix and it seems probable that these are correlated with

developments of the phallosome complex of the male genitalia. Therefore it is

likely that a detailed study of the female genitalia will prove to be of value in

developing a natural classification of the Dixinae.

GENERAL ADULT MORPHOLOGY. There is a remarkable uniformity in

the general external morphology of the adults and very few good taxonomic

characters have been found. The length of the antenna and the shape of the

flagellar segments show the most striking differentiation and form the basis

for the separation of the subfamily into 2 tribes, Dixini and Paradixini. Dif-

ferences in wing venation are present but their usefulness for classification is

limited because of convergence or parallelism in unrelated lines. Color pat-

terns of the body and wings are useful only at the specific level. Secondary

sexual characters of the legs of the male and of the hindtibia may be of some

value in classification but have not been studied extensively.

CHAETOTAXY OF IMMATURE STAGES. Asa result of a comparative

study of the larvae and pupae of species from New Zealand some corrections

have been made in my former interpretation of the chaetotaxy of the Dixinae

(Belkin 1962:94) as indicated in the figures and explained below.

In the pupa, cephalothoracic hair 1-C may be present and it may be possi-

12 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

ble that 2-C is also present but this could not be determined definitely; hair

4-C may be absent. The ventral abdominal hairs have been renumbered and

hair 12 was found on segments II-VI. |

In the larva, the dorsal head hairs have been renumbered. The hair for-

merly interpreted as 0-C belongs to the labrum (palatum) and is eliminated in

the nomenclature while a small hair in the posterior part of the head, not

recognized before, is now interpreted as hair 8-C. Hairs formerly interpreted

as 1,2,3,4,5,6,8-C now become 0,1,3,2,4,5,6-C respectively. On the tho-

rax, hairs 2,3-P,M are interchanged. On abdominal segment II, hair 11 be-

comes 10 and a small hair on the proleg is hair 11; ventral hairs 10-12 on ab-

dominal segments IJI-VII are all renumbered as indicated in the figures.

With this reinterpretation, the chaetotaxy of the Dixinae appears even more

similar to that of the Culicinae, the chief differences being the absence of hair

8 on all abdominal segments of the larva and pupa, the caudal displacement of

hair 8-C on the head capsule of the larva, the absence of hair 12 on abdominal

segments I, II of the larva, and the absence of some hairs on the thorax of the

larva. Even the positions of many hairs follow the same pattern as in the Culi-

cinae.

Almost all hairs in the Dixinae are unbranched, only some of the heavier

larger hairs are barbed and a few of the shorter hairs may have a few branches

in some forms. However, there is considerable differentiation in the length

and thickness of hairs and this character appears to be of considerable impor-

tance in differentiating the 2 tribes and some genera.

Differences in chaetotaxy pattern, including loss of some hairs, have been

noted in the present study. Some of these differences seem to be at a generic

level at least and may prove to be of value in determining evolutionary lines in

the subfamily,

PUPA. The general morphology of the pupa of New Zealand dixa midges

is very uniform but the following characters are apparently of at least generic

value: the length of the antennal sheath, shape of the pinna of the trumpet, and

the chaetotaxy pattern. Specific differences will probably be found primarily in

the spiculation or tuberculation of the integument, in pigmentation, in the shape

of the trumpet and in the length of the paddle. Because of differences in mor-

phology from the Culicinae the measurements of the length of the abdomen and

of the paddle are determined in the following manner. The abdomen is mea-

sured from the anterior border of segment I to the posterior border of segment

VIII; the paddle from the base of segment IX to the apex of the paddle.

LARVA. The general morphology of the larva is quite uniform not only in

the New Zealand species but in all the extralimital forms I have examined.

However, group differences at the tribal and generic level have been noted in

the following: (1) development of "crowns" or rosettes of branched spinules on

the dorsum of abdominal segments, (2) number of abdominal prolegs, (3) ar-

rangement of hooks on prolegs, (4) number of ambulacral combs, (5) spicula-

tion of the antenna, (6) development of tentorium, (7) development of collar on

head capsule, (8) development of the labium and labial process of the head cap-

sule, (9) numerous features of the spiracular apparatus, (10) marginal teeth of

the pecten, (11) sclerotizations of the anal segment, (12) postanal spicules, and

(13) chaetotaxy, including length of hairs. Closely related species appear to be

extremely similar and show differences primarily in pigmentation of the head

capsule, pigmentation and spiculation of the antenna, and spiculation of the

thorax and abdomen.

A special descriptive terminology is used here for the larva of the Dixinae

Belkin: Culicidae of New Zealand 13

primarily for features of the spiracular apparatus and anal segment. The head

length is measured from the anterior border of the labrum (palatum) to a line

connecting the posterolateral angles of the head capsule. The length of the pos-

terolateral lobe is measured in lateral or dorsal aspect from the dorsal end of

the basal unsclerotized line separating it from the pecten plate to the apex of

the lobe exclusive of the marginal spicules. The length of the anal segment is

measured on the dorsal midline from the base of the saddle to the apex of the

distal caudal process. The cylindrical process is the complete sclerotized ring

at the apex of the saddle separated by an unsclerotized line from the distal cau-

dal process.

SYSTEMATICS AND BIOGEOGRAPHY. After this more detailed study of

the dixa midges of New Zealand, particularly of the male genitalia and imma-

ture stages, I am convinced that the similarities between them and true mos-

quitoes are due to common ancestry and not to convergence. It is inconceivable

that the absolute correspondence of all the elements of the chaetotaxy of the

spiracular lobe (or apparatus) and of the 8th (and 9th) abdominal segments of

the larvae of Dixinae and Culicinae, first shown by Martini (1923) and later

confirmed by Belkin (1951), is due to anything other than common ancestry of

the 2 groups. Therefore, I am retaining the dixa midges as the subfamily Dix-

inae of the Culicidae. As before (Belkin 1962:88), I consider that the Dixinae,

Culicinae and Chaoborinae arose contemporaneously from a common ancestor

and that the Dixinae are not in any way ancestral to either of the other 2 sub-

families although they appear to have retained more primitive features than the

latter.

Judging by the morphology of the known existing types, their breeding sites

and their distribution, it seems probable that the Dixinae split into 2 major evo-

lutionary lines very early, probably earlier than the first radiation occurred

in the Culicinae or Chaoborinae, and that neither line has been successful in

occupying a breeding environment different from the original one (ground

waters). This general conservatism is also shown in the preservation of many

primitive general characters in the Dixinae and the great similarity in basic

characters in the larva, pupa and adult within the entire subfamily.

Speciation in the Dixinae appears to have beendetermined largely by unique

specializations in the phallosome of the male genitalia and probably, at least

in some groups, by correlated specializations in the female genitalia, pro-

ducing a mechanical isolating barrier between species. Speciation has been

accompanied by very few and minor changes in the general external characters

of the adults, primarily in the antenna, palpus, wing veins and pigmentation,

legs, and pigmentation of the body. In the immature stages the changes have

also been relatively few and inconspicuous in the general morphology, except in

the trumpet of the pupa and in the spiracular apparatus, anal segment, prolegs

and body spiculation of the larva. The only striking differences noted in the

larvae so far are the development of the abdominal rosettes in the Dixini and the

presence of only 1 pair of prolegs in Meringodixa. However, it is evident from

the present study that obscure but significant differences in the chaetotaxy of

the immature stages distinguish some evolutionary lines in both the larvae and

pupae and that differences insome of the other characters of these stages men-

tioned above are also of taxonomic significance.

It would appear therefore that a natural classification of the subfamily

should be based primarily on fundamental characters of the male and female

genitalia and of the immature stages. These characters must be evaluated

very carefully since parallel developments are evident in the 2 major evolu-

14 Contrib. Amer. Ent. Inst.;- vol 3, noi 1, 1968

tionary lines in the subfamily, e.g. development of a penis filament in the Dix-

ini and Paradixini, elongate flagellar segments in adults of Meringodixa and the

Paradixini, similar type of larval antennal spiculation in Dixini and Paradixini,

and so on.

To date taxonomic studies of Dixinae have consisted largely of species rec-

ognition and diagnosis on the basis of external adult characters and superficial

details of specialization of the sidepiece and clasper of the male. Because the

latter show such excellent diagnostic features that are readily visible, very

little attention has been paid to the complex structures of the phallosome and

proctiger of the male which require dissection and careful interpretation. The

larvae and pupae are undescribed for most species and the few that are known

are not described in sufficient detail to evaluate significant features.

The primary division of the subfamily into the tribes Dixini and Paradixini,

based on characters first recognized by Tonnoir (1924b), appears to be a

natural one and as far as I can determine all the described genera fit into one

or the other readily. The generic classification, on the other hand, is ina

confused state at the present largely because the type species of most genera

have not been described in sufficient detail for the significant taxonomic charac-

ters and stages. The majority of species have been placed in either Dixa or

Paradixa (or Dixella following Lane 1949:257; 1951:336) depending on the length

of the antenna and the shape of the flagellar segments of the adults and some-

times presence or absence of the dorsal abdominal rosettes in the larvae. It is

evident that both genera are polyphyletic assemblages and probably contain spe-

cies which could be placed better into some of the other described genera or

should be segregated into new genera or Subgenera.

The Dixini include the following genera: Dixa Meigen, 1818; Neodixa

Tonnoir, 1925. (= Dixella Tonnoir 1924b); Nothodixa Edwards, 1930; and

Meringodixa Nowell, 1951; the latter was originally placed in a separate mono-

typic tribe because of some annectent characters between the Dixini and Para-

dixini but most of its features warrant its inclusion in the Dixini for the pres-

ent. The Paradixini include 4 genera also: Dixella Dyar and Shannon, 1924;

Dixapuella Dyar and Shannon, 1924; Paradixa Tonnoir, 1924b; and Dixina

Enderlein, 1936. The genus Eriopterites Meunier, 1916 is a homonym of

Erioptera Meigen, 1803 according to art. 56(b) of the International Code of

Zoological Nomenclature since Meunier proposed this name for a fossil species

(tertiaria) which he stated to be without a doubt a species of Erioptera. This

form is not a tipulid but a dixa midge whose generic placement is uncertain.

Therefore no replacement name for Eriopterites is needed at this time.

Details of the affinities and systematics of New Zealand Dixinae are dis-

cussed under each genus and species below. Only the general features of this

fauna are reviewed here. The fauna is harmonious, extensive and entirely

endemic at the species group level. It consists of apparently the most primi-

tive living representatives in each of the 2 major phyletic lines. The Dixini

are represented by the genus Nothodixa (4 sp.) which is known elsewhere only

in Tasmania, New South Wales, Chile and Patagonia, and the derived aberrant

monotypic Neodixa, still known only by the male holotype. Nothodixa probably

dispersed to New Zealand very early at a time when a suitable dispersal route

to South America was also present in the Southern Hemisphere. The New Zea-

land species of Nothodixa have typical disjunct relict populations; there is no

detectable differentiation in the phallosome of the 3 species known in the male

sex or any feature of the pupa in the 2 species known in this stage, and the lar-

vae of the 2 known Species are extremely similar except for modifications inthe

Belkin: Culicidae of New Zealand 15

prolegs. The Paradixini are represented by the genus Paradixa (4 sp.) which

is definitely absent from Chile and Patagonia and whose extralimital distribu-

tion is apparently confined to Tasmania and New South Wales although it is

possible that different phyletic lines assignable to this genus are present in

the Northern Hemisphere. Paradixa probably dispersed to New Zealand con-

temporaneously with most Culicinae after the earlier dispersal of Nothodixa

and at a time when suitable dispersal routes to South America were no longer

available in the Southern Hemisphere. Two of the species (harrisi and

tonnoiri) have typical disjunct relict populations but neozelandica and particu-

larly fuscinervis are relatively widespread and abundant, the latter being one

of the most conspicuous elements of the entire culicid fauna of New Zealand.

All 4 species are very strongly differentiated in male genitalic characters in-

cluding those of the phallosome and proctiger. Only harrisi is strongly dif-

ferentiated in the larva and pupa and has a phallosome markedly different from

the other 3 species and therefore may represent a separate phyletic line from

the latter.

KEYS TO TRIBES AND GENERA

Adults

b, Antenna about twice as long as head and thorax combined; flagellar seg-

ments all cylindrical or filiform, basal one at least 10 times as long

as wide (Paradixini). 5) et SPapadixa

Antenna about as long as head: and thorax combined; ‘flagellar segments

all more or less fusiform, basal one at most 5 times as long as wide

(Dikiniy CONES ee RE eS a atch ae

2(1). Vein R2+3 branched; base of Mg+4 complete. . . . . . Nothodixa

Vein R98 unbranched; base of M3,4 broken (1. minuta) . . . Neodixa

Male Genitalia

1, Penis filament absent; proctiger with sclerotized plates, cercal bristles

2-5, borne on a sclerotized process (Paradixini). . .. . .Paradixa

Penis filament present; proctiger unsclerotized, cercal bristles numer-

ous, borne on a large unsclerotized lobe (Dixini). Nothodixa, Neodixa

Pupae

i Pinna of trumpet narrowly slit all the way basad to apical constriction of

inner tracheoid; antennal case not pei beyond basal third of

wing case (Dixini). . . . Nothodixa

Pinna of trumpet aries slit: ee case ee beyond distal 0.75

of wing case (Paradixini). Be ng tate a were eta eae Dee ae

Larvae

(Neodixa unknown)

1, Dorsum of abdominal segments with conspicuous ovoid rosettes of plu-

mose spinules; none of the hairs of the prothorax reaching anterior

bopaer oc bean Caneule (Nein) ok ak tack os ce cancms ss 4s OLDOGLNA

16 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

Dorsum of abdominal segments without rosettes of plumose spinules;

some of the hairs of the prothorax projecting well beyond anterior

border of head capsule (Paradixini).:2 6). 6540s. Ss Raradixa

TRIBE DIXINI

Genus NEODIXA

1924. Dixella Tonnoir, 1924b:230(4 Dec). TYPE SPECIES: *D. minuta Ton-

noir, 1924b, New Zealand; original designation. Preoccupied by Dixella

Dyar & Shannon, 1924(22 Oct).

1925. Neodixa Tonnoir, 1925:311. TYPE SPECIES: Automatically Dixella mi-

nuta Tonnoir, 1924, New Zealand. Substitute name for Dixella Tonnoir,

1924.

Neodixa of Belkin (1962:103).

Neodixa is characterized by the unbranched vein R9,3. In all other re-

spects the only known specimen (male holotype) fits the diagnosis of Nothodixa

but is different from any species assigned to the latter genus. Edwards (1930:

101) suggested that the venational character of Neodixa might be an individual

variation and considered it hardly necessary to separate it generically, but in

deference to the opinion of Tonnoir he treated it as a distinct genus. I also

prefer to retain Neodixa as a distinct genus with the original characterization

until additional specimens are available for study and it can be determined

whether or not the venational character is genetic.

1. Neodixa minuta (Tonnoir)

Fig. 26

1924. Dixella minuta Tonnoir, 1924b:230. TYPE: *Holotype —, Nelson, N. Z..,

from low shrub along creek, 18 Oct 1923, A.L. Tonnoir [NELS].

Neodixa minuta of Tonnoir (1925:311); Tillyard (1926:352); Edwards (1930a:

101; 1932:15); Taylor (1934:7); Cooper and Rapp (1944:252); Miller (1950:

46); Nowell (1951:241); Belkin (1962:103-104).

FEMALE. Unknown.

MALE: Wing: 2.5mm. Body completely black, head and thorax shining

and with very short bristles, abdomen dull and somewhat velvety. Maxillary

palpus with segment 5 about twice as long as segment 4. Antenna about as long

as head and thorax together; first 7 flagellar segments more or less fusiform

and fairly distinct from one another, segments 8-14 filiform. Legs yellowish

except for tips of femora and tibiae and the ends of the tarsi which are black-

ish; base of hindtarsal segment 1 ventrally with a spinelike bristle; claws pec-

tinate. Wing (Tonnoir 1924b:fig. 8; Belkin 1962:fig.34) dusky and unspotted,

base and anterior border yellowish; vein Sc ending well basad of furcation of R

and somewhat basad of middle of wing; vein R9.3 unbranched and not strongly

arched; base of vein M3g,4 obliterated. Haltere yellowish.

MALE GENITALIA (Tonnoir 1924b:figs.9, 10; Belkin 1962:fig. 33). Similar

Belkin: Culicidae of New Zealand 17

to Nothodixa. Tergite IX with numerous bristles. Sidepiece and clasper sim-

ple. Phallosome apparently similar to Nothodixa. Proctiger simple and with-

out Spines; cercus very prominent and with numerous bristles.

PUPA and LARVA. Unknown.

SYSTEMATICS. The unique known specimen of Neodixa minuta is fully

hardened, not teneral. Except for the aberrant venation it appears to be per-

fectly normal and agrees in general with the characteristics of described spe-

cies of Nothodixa. The 2 aberrant features of the venation, unbranched R9+3

and broken base of Mg,4, are equally well developed on both wings. I have not

seen any such aberrant venation in specimens of Nothodixa but have noted sev-

eral aberrations in the venation, largely unilateral, in specimens of Paradixa

from New Zealand, among them: base of Rg broken (bilateral), apex of Rg

atrophied, apex of Rg fused with R4,5, partial atrophy of Mg,4.

I agree with Edwards (1930a:101) that the holotype of minuta represents a

Species distinct from any other New Zealand dixine. Its characters, other than

the aberrant venation, do not agree with any described species of Nothodixa and

it seems probable therefore that is not a dwarf sport of any of these. I be-

lieve also that the reduced venation of minuta is probably of a genetic nature

and not an individual aberration as suggested by Edwards (loc. cit.). The fact

that venational aberrations of the type present in minuta do occur rarely as in-

dividual variations in some lines does not preclude the genetic fixation of such

an aberrant character ina particular phylad. Since in minuta the unbranched

R2+3 iS a unique exception to the characteristic venation of the entire family

Culicidae in the broad sense, the tendency is to regard this as an individual

variation as Edwards did. However, exceptions to characters of equal impor-

tance, at least at a subfamily level, can occur and are genetically fixed as is

the case in the reduction in the normal number of flagellar segments in the an-

tenna of Ethiopian species of Chaoborus (sen. lat.) notably in Neochaoborus

anomalus (Edwards, 1930). Therefore, the possibility exists that the venational

peculiarities of minuta are genetic and I prefer to regard them as such until

the problem can be finally resolved when additional specimens become available

for study.

BIONOMICS. The specimen was "obtained in sweeping with the net some

low shrubs in a wet part of the track along the 'Reservoir' creek."

DISTRIBUTION (fig.26). NEW ZEALAND. Nelson: Nelson, track along

Reservoir creek, 18 Oct 1923, A.L. Tonnoir, © holotype [NELS].

Genus NOTHODIXA

1930. Dixa (Nothodixa) Edwards, 1930:101. TYPE SPECIES: Dixa campbelli

Alexander, 1922, New Zealand; original designation.

Nothodixa of Belkin (1962:100).

FEMALES (fig.4). Head: Clypeus with numerous bristles distally, some

of these laterally. Antenna about as long as head and thorax together; flagellar

segments 1-5(6) more or less fusiform, segment 1 at most 3 times as long as

maximum width. Thorax: Acrostichals usually extending caudad on prescutel-

lar space; dorsocentrals extending to near parascutellum, posterior part of

row curved laterad; notopleurals numerous; humerals usually few; fossals

numerous; paratergite narrow, bare; parascutellum strongly developed but

18 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

without bristles. Scutellum nearly perpendicular to mesonotum, usually a few

bristles in the middle caudad of apical row; postnotum very large and bulging,

bare. Apn, ppn and upper mep with bristles; stp below transverse suture with

or without short inconspicuous light hairs; apparently no other pleural bristles

or hairs present. Legs: Hindtibia very strongly swollen apically, comb poorly

differentiated, a large ovoid depression just proximad of comb filled with nu-

merous fused spicules which are continued in a narrow line ventrad at base of

comb. Claws of all legs with spicules on outer surface. Wing: Membrane and

veins dusky; microtrichia of membrane large and uniformly densely distributed.

Vein Sc moderately well developed, its apex distinct and ending on C distinctly

basad of furcation of R; crossvein h angled distad apically, a faint crossvein at

level of h connecting Sc to base of R; cross vein r-m always at least its own

length basad of furcation of Rs; crossvein m-cu distinct but weak; bristles on

vein M basad of r-m varied. Alula with a marginal row of dorsal hairs only;

margin of upper and lower calypters bare. Haltere long, winglike; apparently

with 4 rows of bristles, 2 of them marginal. Abdomen: Spiracles present only

on IV-VII. Sternite I without hairs. Genitalia: Segment VIII reduced, its ter-

gite about half as long as that of VII, its sternite subequal to that of VU. Ter-

gite IX small, poorly sclerotized; its lateral lobe short and separated from

median part at least at base. Postgenital plate without distinct median sclero-

tization but with prominant hairy lateral lobes. No distinct bursa copulatrix;

one large spermatheca in segment VII, its duct moderately long, with only 1

loop and with opening on sternite IX.

MALES. Essentially as in the females. Hindtarsal segment 1 with 4 heavy

long spinelike bristles at base on ventral (lower) surface; extreme base of ven-

ter of all tarsal segments 5 with a strong bristle recurved apically. All claws

enlarged and pectinate and with spicules externally.

MALE GENITALIA (fig. 7). Segment VIII: Tergite and sternite at least 0.7

as long as corresponding sclerites of segment VII. Segment IX: Lightly sclero-

tized, without internal ridge articulating with base of sidepiece. Tergite with

bristles, much smaller than sternite, restricted to middle of dorsal surface

and separated from sternite by membrane. Sternite very large, extending to

dorsal surface, bristles restricted to the dorsal lobes. Sidepiece without con-

Spicuous processes but with slight hairy subapical and median basal lobes.

Clasper simple, without differentiated spiniforms. Phallosome: Figured in

everted condition by Belkin (1962, fig. 33) and in normal retracted condition here

(fig. 7); largely membranous. Prosophallus simple, deeply emarginate in the

middle and with a pair of longitudinal sclerites to the aedeagal pouch, widely-

spaced at base. Phallus with small aedeagal pouch which is papillose on inner

surface; aedeagal sclerites forming a poorly sclerotized fused dorsal and dor-

solateral cordate plate from whose thickened apex arises a short free penis fil-

ament within the aedeagal cup; aedeagal sclerites articulating with a long pro-

cess of proximal division of basal piece on each side. Opisthophallus a simple

large transverse lobe with poorly developed, weakly sclerotized lateral scle-

rites articulating with outer part of proximal division of basal piece but not with

the proctiger. Distal division of basal piece strongly developed; proximal di-

vision represented chiefly by inner part (paramere) articulating with aedeagal

sclerite, its outer part very poorly developed. Proctiger largely membranous,

with indistinct basal sclerotizations; cercus a large, prominent lobe with many

bristles.

PUPAE. Cephalothorax: Antennal case not extending beyond basal third of

wing case. Hairs 2,3,4-C apparently all absent; hair 9-C usually far caudad

Belkin: Culicidae of New Zealand 19

near base of wing case, adjacent to hair x-C. Trumpet: Short and broad; pinna

very broad and flaring, narrowly slit all the way basad to apical constriction of

inner tracheoid; apical margin of pinna sinuate, with an evenly spaced row of

simple hooks recurved mesad; external surface tracheoid to near pinna. Meta-

notum: Hairs 11,12-C very close together in a depression; 10-C far mesad of

11,12-C, its distance from the latter several times that between hairs ll and 12.

Abdomen: Hair 1-I-VII far mesad of hairs 2 and 3; hair 2-V-VII far cephalad of

hairs 1 and 3; dorsal sensillum (s) associated with hair 4 on all segments.

Paddle: Hair 1-P at level of base of genital lobe.

LARVAE. Head: Head capsule broad, length about 0.75 of maximum width

or less, strongly emarginate middorsally on caudal border, a more or less

distinct and deep emargination on caudal border ventrolaterally; tentorium very

poorly developed, anterior arm very short, never reaching posterior; labial

process narrow, triangular, pointed distally; collar not developed. Hairs 4,5-C

widely separated, distance between the 2 hairs 4-C subequal to or less than that

between hairs 4 and 5 on one Side; hairs 6,9,10-C near midlevel of head cap-

sule; hairs 12,13-C very close together, their alveoli adjoining; hairs 14, 15-C

both single, 15-C at base of labial process. Antenna: Relatively short, about

0.5 of total head length; relatively straight, without marked curvature in basal

half; shaft with triangular spinules only; hair 1-A inserted near apex, at about

0.8 from base; a clear alveolus (sensillum?) at about 0.7. Thorax: None of

the prothoracic hairs more than 0.5 of entire thorax length, usually much

shorter; hair 2-P at level of 1-P; hairs in group 2, 3-M, T adjoining, separated

from each other by not more than width of 1 basal tubercle. Abdomen: Dorsum

of II-VII with conspicuous oval rosettes of plumose spinules occupying large

part of disc of each segment. Prolegs present on venter of I, II; with heavily

sclerotized, pigmented hooks in 1 rank only on anterior border, other hooks

poorly developed, lightly pigmented and sclerotized, and separated from the

anterior row by a wide space. Ventral ambulacral combs on V, VI only, con-

sisting on each segment of a pair of submedian oval areas with a single rank of

6-8 slightly hooked spines and corresponding internal sclerotized bars; the pair

of oval areas not Separated by median sclerotization; venter of VII with a

prominent median caudal projection with dense spicules. Hair 1-I-V widely

separated from hairs 2,3; dorsal sensillum (s) associated with hair 4 through-

out; hair 14-I laterad of hair 13; hair 11-II within the circlet of caudal hooks

on proleg; hair 12-VII apparently absent; hair 10,11-VII not reaching middle of

segment VIII, hair 6-VII much shorter than hair 7; hairs 3-5-VIII short, none

reaching apex of spiracular lobe. Spiracular Apparatus: Spiracles small,

widely separated, distance between them more than twice length of one antero-

lateral lobe (without spicules). Anteromedian lobe with or without small scle-

rotized plate; hairs 4,5-S closer to spiracle than to hair 3. Anterolateral lobe

small, without conspicuous sclerotized median process mesad of spiracle.

Median plate not developed, lateral sclerotizations small and not connected

across midline, this area more or less densely and uniformly covered with

Short stellate spicules. Posterolateral lobe without apical ventral pointed pro-

cess; hair 8-S very close to marginal spicules, hair 12-S oninner margin of

lobe a considerable distance proximad of apex, hair 13-S on lobe, distad of

hair 14. Pecten plate strongly sclerotized, with simple articulated marginal

teeth and without spicules overlying their bases; hair 1-S simple, inserted in

lower half of plate. Anal Segment: Basal saddle strongly developed, bearing

hair 4c-X; hairs 4-X all short, at most subequal to length of posterolateral

lobe of spiracular apparatus; supraanal spicules in several rows; caudal

20 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

process not distinctly separated from the cylindrical process; hairs 1-3-X

distinctly barbed, moderate in length, less than length of 2 abdominal segments.

SYSTEMATICS. The genus Nothodixa, as currently recognized, is a

strongly differentiated group of true dixines that is restricted to the Southern

Hemisphere and is known at present by 2 species in southeast Australia and

Tasmania, 4 species in New Zealand and 4 species in Chile and Patagonia. All

these species obviously belong to an assemblage distinct from the other generic

group taxa currently recognized in the tribe Dixini except Neodixa. However,

each of the above-mentioned areas appears to have I or 2 endemic groups of

Nothodixa, some of which are so strikingly different in male genitalic and lar-

val characters that they may have to be separated as distinct subgenera and

possibly even genera. The relationships of the different regional groups cannot

be determined at this time because of the extremely poor knowledge of the Aus-

tralian and South American species and the still fragmentary knowledge of the

New Zealand species.

In New Zealand the 4 known species fall into 2 distinct groups on the basis

of wing spotting and infuscation. It seems probable that these groups are also

characterized by larval differences. On the other hand, the 2 groups have ex-

tremely similar male genitalia and pupae.

The campbelli group, consisting of campbelli and philpotti, is characterized

by spotted and infuscated wings and probably also by a larva with hooks simi-

larly oriented and subequal in number on prolegs I and IJ. Contrary to an

earlier suggestion (Belkin 1962:99), it appears now that in the campbelli group,

philpotti is clearly distinct morphologically from the nominate species and is

ecologically isolated from the latter. N. campbelli, as currently interpreted,

has a wide distribution encompassing both South Island and North Island and is

the commonest Nothodixa in New Zealand. There is no indication of geograph-

ical differentiation in this species on South Island and the Wellington area of

North Island where adults have been collected. Farther north only larvae are

known at present and these show only minor differences from those occurring

on South Island.

The otagensis group, comprising otagensis and Septentrionalis, is charac-

terized by wings generally lacking conspicuous infuscations and probably also by

a larva with hooks more numerous and longitudinally oriented on proleg I and

less numerous and transversely oriented on proleg II. It is evident that the 2

forms have arisen as a result of geographical differentiation but because of the

fragmentary nature of the available material it is not clear if this differentia-

tion has reached the specific level. For the present they are regarded as dis-

tinct species.

Much work remains to be done on Nothodixa in New Zealand. The imma-

ture stages are unknown for philpotti and otagensis; the male of philpotti is

unknown and so is the female of septentrionalis. Even for the best known spe-

cies, campbelli, only about 50 adults have been collected or reared and none

north of Wellington. Since there is some indication of both geographical and

ecological differentiation in the known species and since the populations tend to

be disjunct and very small, there is a distinct possibility of unrecognized spe-

cies existing at present in New Zealand. The presence of populations cf larvae

of the campbelli group on Little Barrier and Great Barrier islands is evidence

of the ability of at least one species of the genus either to disperse over mod-

erate ocean barriers or to survive rather major environmental upheavals. A

search should be made for similar isolated populations on other islands in the

New Zealand area.

Belkin: Culicidae of New Zealand 21

The apparent low density population structure, the relict disjunct distribu-

tion and the great uniformity of basic structure in all known species, including

the aberrant Neodixa, suggest that the Nothodixa lineage has been in New Zea-

land a very long time, much longer than the Paradixa lineage. This may ac-

count for the dispersal of Nothodixa to South America at an early date by land

connections through or near New Zealand and also for the absence of Paradixa

in South America because of the absence of such connections later.

BIONOMICS. In New Zealand, Nothodixa seems to be much less common

than Paradixa and its breeding sites appear to be more specialized and re-

stricted. All collections of immature stages of Nothodixa to date have been

in small streams in rather narrow steep valleys in hills and mountains and

never in larger streams or in water courses in flat open valleys or plains.

The 2 species whose larvae are definitely associated appear to differ in their

preference for current strength and for light intensity, campbelli occurring in

Slower flowing water and in more open situations and septentrionalis in fast

water in moderate to deep shade. Even in seemingly favorable habitats the

populations of Nothodixa appear to be usually very small.

Nothodixa larvae can usually be recognized in the field by the projection of

the abdominal rosettes above the water surface film. In collecting containers

they tend to rest with the body straight and not bent into the U-shape charac-

teristic of Paradixa larvae.

Keys to Species

Adults

a Wing conspicuously spotted and infuscated along Some veins . . . 2

Wing usually without spots or infuscations, at most with a faint elaud ¢ on

Pee Oi on ena 1h SERIE PE DAR RN SEAN Ep

2(1). Vein R99 with only a marginal infuscation; vein M proximad of m-cu

usually with an elongate dark streak .. . . 2. campbelli

Vein R943 with a large square or roundish blotch extending into cell Rj;

vein M proximad of m-cu with a large restricted quadrate blotch.

.3. philpotti

3(1). Vein M beyond m-cu subequal to vein Mj,9; mesonotum with reddish

brown Ee Ove and blackish median and egaegac:: stripes.

: 4. otagensis

Vein M ‘beyond m-cu “about ‘1; 5c or more of vein “Mylo: mesonotum uni-

formly brown, without distinct darker stripes . . . 5. septentrionalis

Male Genitalia

(2. philpotti not known)

1; Tergite IX with no more than 15 weak bristles; dorsal lobe of sternite

IX with about 6 weak bristles. . .. «+ &. Campbelli

Tergite IX with at least 20 strong bristles; ‘dorsal lobe of sternite IX

with at least 10 strong bristles. . . . 4. otagensis; 5. septentrionalis

Pupae

Insufficiently known for separation, see 2. campbelli and 5. septentrionalis.

Zz Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

Larvae

(3. philpotti and 4. otagensis not known)

1. Proleg I not markedly larger than II and with 20-30 heavy pigmented

hooks in a transverse are as on proleg II, latter with 15-20 similar

hooks; dorsum of thorax and abdominal segment I with heavy long

spicules or spinules clearly visible at 100X .... . . 4. campbelli

Proleg I twice as large as II and with at least 50 heavy pigmented hooks

in a longitudinal arc, proleg II with 20-30 similar hooks ina trans-

verse arc; dorsum of thorax and abdominal segment I with minute fine

spicules invisible al 100X% . . ..... ... . - . »,..0. Septentrionalis

2. Nothodixa campbelli (Alexander)

Figs. 4,5, 25

1922. Dixa campbelli Alexander, 1922a:20-21. TYPE: Holotype “, Charteris

Bay Banks Peninsula, Canterbury, N. Z., 4 Sept 1921, J. W. Campbell

[ALEX].

Nothodixa campbelli of Belkin (1962:99-100).

Dixa (Nothodixa) campbelli of Edwards (1930a:101; 1932:11); Taylor (1934:7);

Nowell (1951:233).

Dixa campbelli of Alexander (1922b:147); Tonnoir (1924b:224-225, 231-232);

Tillyard (1926:352); Martini (1931:199, 203); Cooper and Rapp (1944:249);

Miller (1950:45).

ADULTS (fig.4). Wing: 3.3-3.5 mm. Forefemur: 1.2-1.4 mm. Abdo-

men: about 2.0 mm. As figured here and as described by Belkin (1962:99- 9-100);

with the following diagnostic features. General coloration yellowish marked

with brown; legs yellowish with tips of femora and tibiae and entire tarsi dark.

Thoracic bristles short and weak, light in color. Mesonotum yellowish, witha

broad median dark stripe on anterior half and a pair of broad lateral dark

stripes beginning about halfway between scutal angle and origin of wing and ex-

tending to scutellum. Stp apparently without hairs. Base of wing, including

membrane, veins C, Sc, and R, yellow to level of crossvein h; a brown streak

on M, a large brown blotch centered on r-m but not reaching furcation of R,

and brown infuscations along R2+3, R445, M34 and Cu and its branches.

MALE GENITALIA (fig.4). As figured here and as described by Belkin

(1962:100). Very similar to otagensis and septentrionalis. Weakly sclerotized

and with weak hairs and bristles. Tergite IX with 6-15 weak bristles, mostly

in distal part. Dorsal lobe of sternite IX with 5-7 weak bristles.

PUPA (fig.5). Abdomen: 3.3 mm. Trumpet: 0.2 mm. Paddle: 0.66 mm.

General morphology and chaetotaxy as figured; very similar to septentrionalis

from which it cannot be differentiated at present. Cephalothorax and metanotum

uniformly moderately pigmented; trumpet uniformly moderately pigmented, a

little brighter than cephalothorax; abdomen and paddle uniformly lightly pig-

mented.

LARVA (fig.5). Head: 0.45 mm. Posterolateral Lobe: 0.34 mm. Anal

Segment: 0.32 mm. General morphology and chaetotaxy as figured; diagnos-

tic characters as in the key. Head: Uniformly moderately pigmented a brown-

ish yellow except for blackish clypeal border, antennal tubercle, eye region,

Belkin: Culicidae of New Zealand 23

caudolateral and caudomesal areas, and posterior border. Hair pigmentation

moderate to strong; hair 4-C projecting far cephalad of clypeal border, the 2

hairs 4-C usually distinctly closer together than hairs 4,5-C on one side. An-

tenna uniformly very darkly pigmented, blackish. Thorax: Dorsum of posterior

part of prothorax and all of meso- and metathorax with large heavy spicules,

sometimes with shorter spinules. Hairs moderately to strongly pigmented,

particularly on dorsum; at least hair 4-P longer than head capsule. Abdomen:

Dorsum of segment I with spicules or spinules similar to those on dorsum of

thorax. Dorsal hairs of segment I strongly pigmented and heavy as on thorax.

Disc within the rosettes of segments II-VII with heavy, darkly pigmented, broad,

apically-branched spicules with distinct sclerotized basal area; distance be-

tween bases of spicules mostly subequal to length of spicules, rarely more than

twice their length. Proleg I only slightly larger than proleg II, with 20-30 pig-

mented heavy hooks; proleg II with 15-20 similar hooks. Hairs on all segments

moderately pigmented and moderately long. Spiracular Apparatus: Anterome-

dian lobe without sclerotization; hair 13-S very heavy, usually nearly straight;

pecten teeth usually heavy. Anal Segment: Hair 4b-X about as thick as hair 4-C

and usually longer than latter. ,

SYSTEMATICS. N. campbelli seems to be the only widespread and the

commonest species of Nothodixa in New Zealand. I have examined adults from

all the land districts of South Island except Marlborough and Southland, and

also from the vicinity of Wellington. In this larger series of adults the same

wide range of coloration of the body and infuscation of the wing was found as

was noted earlier (Belkin 1962:100). This variation is neither geographical nor

ecological, the whole range occurring in different localities and in different

individuals reared from the same breeding site. Perhaps most of the variation

is due to differences in the stage of hardening or to the age of the individuals.

I am convinced that the adults I have seen represent only one species and that

there is no geographical subspecific differentiation within the range encom-

passed by these specimens.

I have included provisionally under campbelli all larvae of Nothodixa with

strong spiculation of the thoracic dorsum and with the hooks similarly oriented

and subequal in number on prolegs I and II. There is considerable variation in

several characters in these larvae and the possibility exists that 2 or more

Species are represented in this sample. Only through association of stages by

individual larval rearings can the problem be resolved.

The unassociated whole larvae of campbelli from the type locality and all

the collections on South Island except Black Gully (Otago) conform quite well to

the description given above which is based on larval skins associated with in-

dividual adults. There is some variation in the distance between head hairs

4-C, in the length of prothoracic hairs, and in the length and spacing of the

dorsal thoracic spicules. The latter may be due primarily to unequal wear of

the spicules and to differences in the stretching of the integument in larvae of

different sizes.

The larvae from Black Gully (Otago), the southernmost reported locality

for Nothodixa, at first glance appear to be strikingly different from typical

campbelli in the much shorter and more widely spaced dorsal thoracic spic-

ules, the space between the bases of the spicules being usually 3 times or more

the length of anindividual spicule. In addition the thoracic and abdominal hairs

are usually considerably shorter, anal hairs 4-X are shorter and weaker, and

the spicules within the abdominal rosettes are shorter and more widely spaced.

There is considerable variation in all these characters in the small series of

24 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

larvae (9) but there does not seem to be any significant overlap with typical

campbelli. Furthermore, hair 13-S on the posterolateral lobe of the spiracu-

lar apparatus is considerably longer than in typical campbelli and is apically

recurved, a condition never encountered in the latter. It is possible that all of

these differences are individual environmental modifications induced by the

swifter current in which these larvae were found, and that therefore we are

dealing here with an extreme ecotype of campbelli. I suspect, however, that

this is a different species, either an undescribed one or philpotti which proba-

bly breeds in swifter streams than campbelli and should havea larva in general

Similar to the latter. Had these larvae been found in a locality where philpotti

is known, I would not hesitate to assign them provisionally to that species.

The 9 larvae of the campbelli type known from 6 different localities on

North Island, 2 of them in association with septentrionalis, also differ from

typical campbelli but in fewer features than the larvae from Black Gully. The

chief differences are in the shorter and more widely spaced dorsal thoracic

spicules (longer than in Black Gully population), smaller spicules within the

rosettes, and Somewhat longer hair 13-S, which however is nearly straight.

These differences are so minor that it seems unlikely that a species different

from campbelli exists on NorthIsland. However, this possibility should not be

dismissed and an effort should be made to rear material from the North Island

populations to resolve the problem.

BIONOMICS. The immature stages of campbelli have been found most fre-

quently along the margins of very small tributaries of streams in rather nar-

row steep valleys, never along the margins of larger streams or in flat open

valleys. In several instances the water courses used by campbelli have been

mere trickles or even small residual pools without detectable flow. The imma-

ture stages have been found also along the grassy margins of small streams

with slow to moderate current but no specific information was obtained as to

the strength of the current in the exact location where the larvae were found.

Concentration of larvae and pupae have been found in flotage blocking the flow

of water. The general impression is that this species prefers quiet water and

that it can utilize quiet pools or marginal pockets in streams with considerable

current. On South Island campbelli is frequently found associated with Paradixa

fuscinervis but the latter has a much wider range of breeding sites. On North

Island campbelli has been found together with septentrionalis in 2 collections

but it is probable that campbelli was utilizing quieter water.

Adults of campbelli have been collected in small swarms over a stream

(Alexander 1922a:21).

DISTRIBUTION (fig.25). Material examined: 531 specimens; 14 0; 37 9;

422 larvae; 58 pupae; 31 individual rearings (20 larval, 3 pupal, 8 incomplete).

NEW ZEALAND. North Auckland: Mangamuka Gorge, 2 mi N of Soda |

Spring, 21 Dec 1963, BS (NZ 41), 11(41-102) [UCLA]; elev. ca 500 m, 18 Feb

1965, F (NZ 205), 2 L[UCLA].

Little Barrier Island: Stream between Te Waikohare and Tirikikawa, elev.

7m, 6 Mar 1964, S(NZ 128), 1 L[UCLA].

Great Barrier Island: Port Fitzroy, 12 Mar 1964, S (NZ 138), 1 L [UCLA].

South Auckland: Te Aroha, Stoney Creek, 26 Mar 1964, S(NZ 139), 2L

[UCLA]. |

Gisborne: Urewera National Park, Hopuruahine area, Orangihikoa stream,

elev. 2450 ft, 30 Mar 1964, S (NZ 141), 2 L[UCLA].

Wellington: Wilton Bush (as Welton's Bush), Mar, A.L. Tonnoir, A [Ton-

noir 1924b]. Day's Bay, 29 Nov 1921, A.L. Tonnoir, 1 A [NELS].

Belkin: Culicidae of New Zealand 25

Nelson: North Branch Riwaka River, 4 Jan 1964, BG (NZ 75), 1 1p (75-

101), 3 1P (75-102-104), 23 L, 29[UCLA]. Pokororo, 19 Dec 1921, A.L. Ton-

noir, 1 A[NELS]. The Glen, 12 Mar 1964, S (NZ 168), 1 1p? (168-103), 1 po’

(168-104), 10 L, 4 P[UCLA]. Nelson, junction of Maitai River and Sclanders

stream, 11 Feb 1964, S (NZ 135), 2 L [UCLA]; South Branch Maitai River, 11

Feb 1964, S(NZ 127), 1 lpo (127-102), 11P (127-104), 11 L[UCLA]. Nelson,

8Apr 1922, A.L. Tonnoir, 2 A|NELS], 1?[CANT]; 27 June 1922, A.L. Ton-

noir, 12[CANT]; 10 Aug 1922, A.L. Tonnoir, 1 A[NELS]; 18 Oct 1923, A. L.

Tonnoir, 1 o [CANT]. Dodson's Valley, elev. ca 68m, 12 Mar 1964, S (NZ -

165), 2 1p? (165-101, 103), 1 1po’ (165-105), 21P (165-102, 104), 85 L, 6 P, 19

[UCLA]. Aniseed Valley, 5 Jan 1964, BG (NZ 79,80), 10 1p? (79-101-107; 80-

102-104), 75 L, 8P, 10, 49(79), 40 L, 2 P (80) [UCLA]. New Creek Rd,

between Murchison and Kawatiri Junction, 8 Feb 1964, S (NZ 116), 1 L [UCLA].

Westland: Lake Brunner, Feb, A.L. Tonnoir, L[Tonnoir 1924b]. Kumara,

15 Sept 1929, J.W. Campbell, 19 [USNM], 12 [UCLA]; 6 Nov 1929, J. W.

Campbell, 40°, 1 2[USNM], 1@[UCLA]. Waiho, 27 Jan 1922, A.L. Tonnoir,

1, 19[NELS].

Canterbury: Cass, 30 Nov 1924, A.L. Tonnoir, 1 “ [CANT]. Christchurch,

July, A.L. Tonnoir, A [Tonnoir 1924b]. Governors Bay, 27 Sept 1923, J. F.

Tapley, 1% |BMNH]; 3 Jan 1964, BD (NZ 72), 3 1p? (72-102, 103,109), 11P

(72-105), 80 L, 2 P[UCLA]. Charteris Bay, 4 Sept 1921, J.W. Campbell, &

holotype [ALEX, not seen], 6 o paratypes[CANT]. Hilltop, 14 Jan 1925, A.L.

Tonnoir, 1 2 [CANT].

Otago: Ben Lomond, 2 Feb 1964, S(NZ 111), 1 po (111-101), 1 p? (111-

103), 51 L, 6 P[UCLA]. Black Gully, 31 Jan 1964, S(NZ 109), 9 L[UCLA].

3. Nothodixa philpotti (Tonnoir)

. Fig. 25

1924. Dixa philpotti Tonnoir 1924b:227-228. TYPE: *Holotype $, Nelson,

South Island, N. Z., sweeping along mountain creek, 5 Dec 1921, A.L.

Tonnoir [NELS].

Nothodixa philpotti of Belkin (1962:100-101).

Dixa (Nothodixa) philpotti of Edwards (1932:11); Taylor (1934:7); Nowell (1951:

239).

Dixa philpotti of Cooper and Rapp (1944:251); Miller (1950:46).

FEMALE (Tonnoir 1924b:fig.3; Belkin 1962:fig. 34). Wing: 3.5 mm. Fore-

femur: 1.5mm. Abdomen: about 2.5 mm. In general similar to campbelli

but darker in coloration and with the following diagnostic features. Tho-

racic bristles dark; light coloration of mesonotum yellowish orange, brown

stripes more extensive than in campbelli; pleuron largely dark, yellowish orange

along mesonotal border and above hindcoxae. Femora infuscated in the middle

and with a subterminal ring of distinctly lighter coloration; tibiae and tarsi

entirely dark. Base of wing dusky, only slightly lighter than rest; blotch on

r-m large, extending all the way to furcation of R; vein M proximad of m-cu

with a large quadrate blotch extending well into cell R; vein Ro+3 with a large

quadrate or roundish blotch extending well into cell Ry; veins R4,5, M3+4, Cu

and its branches, and anal margin of wing with broad lines of infuscation; vein

Sc strong but relatively shorter, ending on C before 0.7 of distance between

26 Contrib. Amer. Emt. nst., vol. 3, uo. 1, 1968

crossvein h and furcation of R, with bristles on distal part; vein Rs beyond

r-m about 1.5 of r-m length; vein Ro9,3 very strongly arched at base, usually

with a stump vein in blotch in middle of curvature; distal part of Cu, strongly

recurved. Haltere usually infuscated on stem.

MALE and PUPA. Unknown.

LARVA. Unknown but probably of the type of the Black Gully (Otago) pop-

ulation briefly described in the section on SYSTEMATICS under campbelli,

which may possibly belong to philpotti.

SYSTEMATICS. In general morphology and particularly wing infuscation,

philpotti comes closest to campbelli and I consider that these 2 forms consti-

tute a distinct group. I am convinced that philpotti is a species distinct from

campbelli since in the additional material of both forms that I have examined

there is no indication of intergradation. Of the 5 specimens of philpotti that I

have seen only the one in BMNH lacks the stump vein on R93. N. philpotti is

probably a derivative of the campbelli group adapted for breeding in fast

flowing water.

BIONOMICS. This species probably breeds in fast flowing mountain

streams as all known adults have been ''collected by Sweeping the plants along

mountain creeks with a much swifter flow of water than in those the Dixa lar-

vae usually breed" (Tonnoir 1924b:228). The larvae from Black Gully (Otago)

mentioned above were apparently collected in such a situation and may possi-

bly belong to this species.

DISTRIBUTION (fig. 26). Material examined: 5 9.

NEW ZEALAND. Nelson: Nelson, 18 Oct 1923, A.L. Tonnoir, 1 ¢ para-

type [CANT]; 5 Dec 1921 (not 15 Dec), A.L. Tonnoir, ? holotype [NELS]; 15

Dec 1921, A.L. Tonnoir, 1 2? paratype [NELS], 1 2 paratype |[BMNH]. Mt. Ar-

thur, elev. 4000 ft, 24 Dec 1921, A.L. Tonnoir, 12[NELS]. This species

has not been found definitely in any other locality but it is possible that it has

a wider distribution since larvae from Black Gully (Otago) listed and discussed

under campbelli may actually belong to philpotti.

4. Nothodixa otagensis (Alexander)

Figs. 7, 26

1922. Dixa otagensis Alexander, 1922b:147. TYPE: Holotype 3, Ben Lomond,

Otago, N. Z., 30 Dec 1921, G. Howes [ALEX].

_Nothodixa otagensis of Belkin (1962:101-102).

Dixa (Nothodixa) otagensis of Edwards (1932:11); Taylor (1934:7); Nowell (1951:

239).

Dixa otagensis of Tonnoir (1924b:225-226); Cooper and Rapp (1944:251); Miller

(1950:45).

ADULTS (Tonnoir 1924b:fig.11; Belkin 1962:figs. 32,34). Wing: 5.0 mm.

Forefemur: 2.0 mm. Abdomen: about 2.5 mm. As described by Belkin (1962:

101); in general similar to campbelli but wing without conspicuous infuscations

and with the following diagnostic features. General coloration brown to red-

dish brown. Thoracic bristles moderately long, all dark except those on stp;

ground color of mesonotum reddish brown, with broad median and posterolat-

eral longitudinal stripes. Wing membrane without infuscations except for a

faint cloud on r-m (indistinct or apparently completely absentin dry specimens);

Belkin: Culicidae of New Zealand 27

furcation of R943 at level of middle of Cuy; vein M beyond m-cu at most equal

to Mj,9; furcation of M distinctly basad of level of end of Cuy.

MALE GENITALIA (fig. 7). As figured; in general very similar to camp-

belli and septentrionalis. Moderately sclerotized and with strong hairs and

bristles. Tergite IX with 26-28 bristles, more or less evenly distributed. Dor-

sal lobe of sternite IX with 12, 13 bristles.

PUPA and LARVA. Unknown; probably of the septentrionalis type.

SYSTEMATICS. After examination of additional material, I agree with

Tonnoir (1924b:227) that otagensis and septentrionalis should be treated as dis-

tinct but closely related species and I am recognizing the otagensis group for

these 2 species. Our present knowledge of both species is still very inadequate

and fragmentary and it is possible, as originally suggested by Tonnoir (loc.

cit.), that the 2 forms are merely geographical races of 1 species. Whether or

not this is the case can be determined only when material of the group becomes

available from the area between the widely disjunct known distributions of the

2 forms.

The differences in size, color of thorax and in wing venation mentioned in

the diagnosis are quite distinct. In otagensis there is little individual variation

in these characters except for 1 specimen which shows the extreme condition

in the shortness of R9,3 and M beyond m-cu illustrated by Tonnoir (1924b:fig.

11), and which was marked by him with a manuscript name (see DISTRIBUTION

below). This type of variation in the southernmost locality of otagensis may

have suggested to Tonnoir that farther north there might be the reverse trend

toward the condition in septentrionalis in wing venation as well as in other

characters.

Whatever the taxonomic status of otagensis and septentrionalis actually is,

it is evident that in the otagensis group there has been geographical differenti-

ation within a single lineage. This appears to be the only clear-cut instance of

this phenomenon in the Culicidae of New Zealand.

BIONOMICS. Nothing is known of the bionomics of otagensis; even the cir-

cumstances of the collections of adults are not recorded. If otagensis and

septentrionalis are truly closely related, it is very likely that the immature

stages of otagensis will be found in fast flowing small streams similar to those

used by septentrionalis.

DISTRIBUTION (fig. 26). Material examined: 8 specimens; 50, 29, 1A

sex undetermined.

NEW ZEALAND. Westland: Waiho, 28 Jan 1922, A.L. Tonnoir, 1 A [NELS].

Canterbury: Christchurch, Jan 1922, J.W. Campbell, 1c, 1? [BMNH].

Otago: Glenorchy, 3 Jan 1923, C.C. Fenwick, 1c [BMNH]. Ben Lomond,

30 Dec 1921, G. Howes, & holotype, 1 A [ALEX, not seen]. Queenstown, 1

Feb 1922, A.L. Tonnoir, 2 o (1 marked type ''meridionalis"), 19 [NELS], 1¢

[CANT].

5. Nothodixa septentrionalis (Tonnoir)

_ Figs. 6, 7, 26

1924. Dixa septentrionalis Tonnoir 1924b:226-227. TYPE: *Holotype o, Te

Aroha, South Auckland, N. Z., from rock at foot of waterfall, 28 Feb

1923, A.L. Tonnoir [NELS].

28 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

Nothodixa septentrionalis of Belkin (1962:102). |

Dixa (Nothodixa) septentrionalis of Edwards (1932:11); Taylor (1934:7); Nowell

(1951:239).

Dixa septentrionalis of Cooper and Rapp (1944:251); Miller (1950:46).

FEMALE. Unknown.

MALE (Tonnoir 1924b:fig.5; Belkin 1962:fig.34). Wing: 3.5mm. Fore-

femur: 1.5 mm. Abdomen: about 2.5 mm. As described by Belkin (1962:

102); similar to otagensis but differing from it in the following diagnostic fea-

tures. General color uniformly brown, mesonotum without distinct dark longi-

tudinal stripes. Thoracic bristles stronger than in otagensis. Wing membrane

entirely without infuscations, no cloud on r-m; furcation of Ro,9 at level of

apex of Cu, or beyond; vein M beyond m-cu at least 1.5 of Mj.9; furcation of

M distad Q: level of apex of Cuy.

MALE GENITALIA (fig. 7). As figured; in general very similar to camp-

belli and otagensis. Lightly sclerotized but with strong hairs and bristles. Ter-

gite IX with 20-21 bristles, more or less evenly distributed. Dorsal lobe of

sternite IX with 10,11 bristles.

PUPA (fig.6). Abdomen: 3.5mm. Trumpet: 0.3 mm. Paddle: 0.66 mm.

General morphology and chaetotaxy as figured; very similar to campbelli from

which it cannot be differentiated at present. Cephalothorax and metanotum

uniformly moderately pigmented; trumpet uniformly moderately pigmented; a

little brighter than cephalothorax; abdomen and paddle uniformly lightly pig-

mented.

LARVA (fig.6). Head: 0.46 mm. Posterolateral Lobe: 0.34 mm. Anal

Segment: 0.32 mm. General morphology and chaetotaxy as figured; diagnos-

tic characters as in the key; in general similar to campbelli from which it dif-

fers chiefly in the following. Head: Uniformly lightly pigmented a clear yellow

except for same darkened areas. Hairs shorter, more slender and usually

lighter in color; hair 4-C barely projecting cephalad of clypeal border, the

distance between the 2 hairs 4-C usually subequal to or greater than between

4,5-C on one side. Thorax: Dorsum with numerous minute spicules arranged

in short transverse rows, practically invisible at 100 X. Hairs shorter and

lighter in coloration; all hairs, including 4-P, shorter than head capsule. Ab-

domen: Dorsum of segment I with minute spicules as on thorax. Dorsal hairs

of segment I lightly pigmented and short as on thorax. Disc within the rosettes

of segments II-VI with much smaller, lightly pigmented, broad, apically-

branched spicules without distinct sclerotized basal area; distance between

bases of spicules at least 3 times length of spicules. Proleg I more than twice

as large as proleg II, with 50 or more pigmented hooks arranged in a longitu-

dinal arc; proleg II with 20-30 hooks arranged in a transversearc. Hairs on all

segments shorter and lighter in color. Spiracular Apparatus: Anteromedian

lobe with distinct small sclerotization, bearing hair 3-S on each side; hair 13-S

longer and not as heavy, its distal part distinctly recurved; pecten teeth usual-

ly smaller. Anal Segment: Hair 4b-X much thinner and usually shorter than

hair 4c. -

SYSTEMATICS. N. septentrionalis differs from otagensis chiefly in size, —

color and wing venation. It is treated here as a distinct species but it may

possibly be only a geographical race of otagensis (see). However, in the small

series of adults (7) there is no overlap of diagnostic characters with the latter

species.

BIONOMICS. The immature stages of septentrionalis have been found in

Belkin: Culicidae of New Zealand 29

small mountain or hill streams with moderate to strong current. This species

seems to prefer moderate to deep shade and the margins of streams with strong

current. In collecting containers, the larvae rest more frequently with the

body fully extended than in the U-bend characteristic of most dixids. As indi-

cated under campbelli, 2 collections of septentrionalis included a few larvae of

the former but it seems probable that these were collected in quieter water.

The male adults of the type series were all collected on rocks at the foot

of a waterfall.

DISTRIBUTION (fig. 26). Material examined: 84 specimens; 7, 3 pupae,

74 larvae; 2 larval individual rearings (1 incomplete).

NEW ZEALAND. South Auckland: Te Aroha, 28 Feb 1923, A.L. Tonnoir,

So holotype, 3 0 paratypes [Nels], 1 o paratype [CANT, I- 292], & paratype

[BMNH]. Stoney creek, 26 Mar 1964, S (NZ 139), 1 lpo’ (139-101), 11P (139-

102), 50 L [UCLA].

Gisborne: Urewera National Park, Hopuruahine area, elev. ca 2900 ft, 27

Dec 1963, BS(NZ 56), 2 L [UCLA]; Hopuruahine area, Orangihikoa stream,

elev. 2450 ft, 30 Mar 1964, S (NZ 141), 20 L, 1 P[UCLA].

TRIBE PARADIXINI

Genus PARADIXA

1924. Dixa (Paradixa) Tonnoir, 1924b:222. TYPE SPECIES: *Dixa neozelandica

Tonnoir, 1924, New Zealand; selection of Edwards 1932:9.

Paradixa of Belkin (1962:105-106).

Dixella in part of Lane (1951); Nowell (1963); Hubert (1965); Peters and Cook

(1966).

FEMALES (figs.1,8). Head: Clypeus with a few bristles apically, in the

middle only. Antenna at least 1.5 as long as head and thorax together; flagellar

segments all elongate, cylindrical or filiform, segment 1 at least 8 times as

long as wide. Thorax: Acrostichals usually extending caudad on prescutellar

space; dorsocentrals extending to near parascutellum, posterior part of row

curved laterad; notopleurals usually 2-3; humerals usually numerous; fossals

1-2; paratergite narrow, bare; parascutellum strongly developed but without

bristles. Scutellum nearly perpendicular to mesonotum, without bristles caudad

of apical row; postnotum very large and bulging, bare. ‘Apn and ppn with a few

bristles, stp without hairs; apparently no other pleural bristles or hairs pres-

ent. Legs: Hindtibia moderately swollenapically, comb strongly differentiated,

a narrow longitudinal ridge just proximad of comb, apparently consisting of

fused spicules which are continued in a narrow line ventrad at base of comb.

Claws of all legs with spicules on outer surface. Wing: Membrane and veins

dusky; microtichia of membrane large and uniformly densely distributed. Vein

Sc usually strongly developed and with bristles (except in tonnoiri), its apex

ending on C at or near level of furcation of R; cross vein h angled distad apical-

ly, a faint crossview at level of h connecting Sc to base of R; crossvein r- r-m at

or a Short distance (less than own length) distad of furoation of Rs; crossvein

m-cu very weak, sometimes indistinct; bristles on vein M basad of r-m varied.

Alula with a marginal row of dorsal hairs only; margin of upper and lower

calypters bare. Haltere long, winglike; apparently with 4,5 rows of bristles,

30 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

2 of them marginal. Abdomen: Spiracles present only on IV-VII. Sternite I

without hairs. Genitalia: Segment VIII large, its tergite subequal to that of VII,

its sternite enlarged, about twice as long as that of VII. Tergite IX large, dor-

sal part strongly or weakly sclerotized; its lateral lobe joined to median part

at least at base. Postgenital plate strongly sclerotized in middle and projecting

as a triangular process beyond the small hairy lateral lobes. A distinct bursa

copulatrix present in all species, with strong internal sclerotizations; 1 large

spermatheca in segment VII, its duct varied in length and with opening in dor-

sal wall of bursa.

MALES. Essentially as in the females. Hindtarsal segment I without

differentiated spinelike bristles at base on ventral (lower) surface; tarsal

segments 5 without specialized recurved bristle at base. All claws enlarged and

with spicules externally; those of foreleg and midleg pectinate; those of hind-

leg simple or with 1 tooth only.

MALE GENITALIA (fig. 7). Segment VIII: Tergite and sternite about 0.5

of corresponding sclerites of segment VII. Segment IX: More or less strongly

sclerotized and forming a complete ring. No membrane separating tergite

from sternite; boundary of tergite indicated by internal ridge on each side far

ventrad which provides a sternal articulation for sidepiece. Sternite thus re-

stricted to ventral surface, with bristles. Tergite with or without median dor-

sal caudal lobe, with bristles. Sidepiece with or without conspicuous lobes.

Clasper irregular in shape, with moreor less distinct processes or lobes; sev-

eral to many spiniforms present. Phallosome: Very complex and with strong

sclerotizations. Prosophallus strongly developed and usually heavily sclero-

tized (except in harrisi); formed by a prosophallic sclerite continuous across

midline sternally but usually interrupted by a membrane on midline of tergal

surface (except in harrisi in which it encloses an unsclerotized median area).

Phallus with large broad shallow aedeagal pouch; penis filament not developed;

aedeagal sclerites broad, not distinctly separated from central part of proxi-

mal division of basal piece; separated from each other on midline by mem-

brane and not joined tergally. Opisthophallus very strongly developed; lateral

sclerite articulated with outer angle of outer part of the proximal division of

basal piece and with the base of the proctiger; with projecting spines or pro-

cesses. Distal division of basal piece moderately developed; proximal divi-

Sion very strongly developed, with both a central part continuous with aedeagal

sclerites anda strongly sclerotized outer part articulating with opisthophallus.

Proctiger very prominent and complex, articulating at base with opisthophallus;

heavily sclerotized at base and bearing 1 or more spinelike processes with

heavy apical spiniforms; median portion, bearing anus, membranous; cercus

represented by more or less prominent lobe bearing 2 or more Simple setae.

PUPAE. Cephalothorax: Antennal case extending beyond distal 0.75 of

wing case. Hair 2-C apparently always absent; hair 1-C apparently sometimes

absent; hair 3-C always well developed; hair 4-C always present; hair 9-C

usually near 8-C not x-C. Trumpet: short and broad; pinna narrow or broad-

ened but always without slit; apical margin of pinna smooth or sinuate, with

evenly-spaced row of rather broad crenulations hooked mesad; external surface

tracheoid at base only. Metanotum: Hairs 11, 12-C very close together in a de-

pression; position of hair 10-C varied. Abdomen: Hair 1-I-VI near hairs 2 and

3; hair 2-V-VIII near level of hairs 1 and 3; dorsal sensillum(s) associated with

hair 3 on all segments. Paddle: Hair 1-P at level of base of genital lobe or

distad.

LARVAE. Head: Head capsule relatively longer than in Nothodixa, its

Belkin: Culicidae of New Zealand | 31

length about 0.9 of maximum width or more, less strongly emarginate middor-

sally on caudal border, nodistinct emargination on caudal border ventrolateral-

ly; tentorium strongly developed, anterior arm very long, reaching posterior;

labial process broad, truncate or rounded distally; collar more or less strongly

developed. Hairs 4, 5-C close together, distance between the 2 hairs 4-C twice

or more that between hairs 4 and 5 on one Side; hairs 6,9, 10-C in posterior half

of head capsule; hairs 12, 13-C separated from each other by several times width

of their alveoli; hairs 14,15-C both branched, 15-C near middle of labial pro-

cess. Antenna: Relatively long, about 0.6 or more of total head length; dis-

tinctly concave mesad in basal half; ventral surface with short triangular spin-

ules replaced by hairlike spicules distad of hair 1-A toward inner margin; hair

1-A inserted in basal0.67; subapical clear alveolus apparently absent. Thorax:

Some of the prothoracic hairs as long as entire thorax; hair 2-P near level of

3-P; hairs in groups 2, 3-M, T widely separated, distance between them several

times width of 1 basal tubercle. Abdomen: Dorsum of segments without rosettes

of plumose spinules. Prolegs present on venter of I, II; with heavily sclerotized

pigmented hooks in 2 ranks, other small lighter hooks immediately adjacent

caudad. Ventral ambulacral combs on V-VIII consisting on each segment of a

pair of submedian straight lines of 6-8 slightly hooked spines in1 rank and cor-

responding internal sclerotized bars; the pair of lines separated by a small

median sclerite; venter of VII without prominent median caudal projection. Hair

1-I-V widely separated from hairs 2,3; dorsal sensillum(s) associated with

hairs 2, 3 throughout; hair 14-I mesad of hair 13; hair 11-II caudad of hooks on

proleg; hair 12-VII present; at least hair 11-VII extending beyond middle of

segment VIII, hair 6-VII at least 0.67 of hair 7; hairs 3-5-VIII long, at least 1

of these reaching apex of spiracular lobe. Spiracular Apparatus: Spiracles

relatively larger than in Nothodixa and less widely spaced, distance between

them not more than1.5 length of 1 anterolateral lobe (without spicules). Antero-

median lobe with a forked basal sclerotization; hair 3-S on the forks, hairs

4,5-S closer to hair 3 thanto spiracle. Anterolateral lobe moderately large,

with a conspicuous sclerotized median process mesad of spiracle. Median plate

with 2 paired and 1 unpaired median plates developed; the paired ones continu-

ous with lateral plates, all plates with sharp simple spinules; membranous area

between them with thin hairlike spicules. Posterolateral lobe with apical ven-

tral pointed process; hair 8-S removed ventrad from lateral spicules, hair 12-

S near apex of lobe, hair 13-S not on lobe proper, basad of hair 14. Pecten

plate usually poorly sclerotized, with complex branched marginal spicules

or spinules and with smaller complex spicules overlying their bases; hair 1-S

always forked or branched, inserted near the middle of plate or in upper half.

Anal Segment: Basal saddle moderately developed, not bearing any hairs; hairs

_ 4-X all much longer than posterolateral lobe of spiracular apparatus; supra-

anal spicules in 1 rank only; caudal process more or less distinctly separated

from cylindrical process; hairs 1-3-X without conspicuous barbs, longer than

combined length of 2 abdominal segments.

SYSTEMATICS. I am not accepting the synonymy of Paradixa with Dixella

Dyar and Shannon, 1924 (22 Oct; type species D. lirio Dyar and Shannon, 1924,

from Panama) which was first proposed by Lane (implied in 1949:257; stated

in 1951:336) and which has been followed by Nowell (1962), Hubert (1965), and

Peters and Cook (1966), apparently in every case without examining in detail

the respective type species. Only 2 other paradixine generic group taxa have

been proposed, both monotypic, Dixapuella Dyar and Shannon, 1924 (D. mar-

ginata Loew, 1863, Washington, D.C.), and Dixina Enderlein, 1963 (D. ob-

32 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

scura Loew 1849, Europe). No serious attempt at a classification of the para-

dixines has ever been made and the tendency has been to describe species pri-

marily on the basis of wing color pattern and venation and superficial structures

of the male genitalia (clasper and sidepiece specializations), and to lump them

all into 1 genus. Peters and Cook (1966) described what I believe to be the sig-

nificant features of the phallosome and proctiger of the males but did not fig-

ure them or use them to group species. It is evident that on the basis of these

structures at least 3 distinct major phylads were included in their concept of

Dixella, possibly one of them congeneric with lirio and another with neozeland-

ica. This view is supported by differences I have noted in the larvae of a few

species from California.

For the present I am characterizing Paradixa primarily on the features

of the male phallosome and proctiger listed above in the diagnosis of the genus.

In addition to the 4 New Zealand species treated here the genus includes 2 spe-

cies from Tasmania (tasmaniensis and unipunctata) and 2 species from New

South Wales (nicholsoni and humeralis) all described by Tonnoir (1924a). No

other species can be definitely assigned to Paradixa at this time but it appears

that this genus may be represented by several forms in the Holarctic region. No

Species of Paradixa nor any paradixineat all is known from Patagonia and South

Chile. None of the "neotropical" species of Dixa(Dixella) of Lane (1953) appears

to be a true Paradixa.

The New Zealand species of Paradixa fall into 2 groups on the basis of

characters of the male genitalia, larvae and pupae. The single known species

in the harrisi group appears to have a relict population structure and is cur-

rently known only on North Island. The neozelandica group includes 3 species,

one of which, tonnoiri, appears to have a relict population structure and is

known only on South Island. The other 2 species of the group, neozelandica and

fuscinervis, are dominant modern forms with nearly identical wide distribu-

tions throughout both North Island and South Island. The 3 species of the neo-

zelandica group are obviously derivatives of the same stock; their larvae and

pupae are very similar and the only striking differences are in unique special-

izations of the male genitalia. Each species has a distinct wing pattern but the

differences are primarily in the extent and density of infuscation.

There is no gross ecological differentiation in species of Paradixa in New

Zealand, the 4 species usually occurring sympatrically, even in the same

breeding site, tonnoiri, neozelandica and fuscinervis on South Island, and

harrisi, neozelandica, and fuscinervis on NorthIsland. There is no indication

of geographical differentiation in the 2 dominant modern species while the allo-

patric distributions of tonnoiri and harrisi involve members of 2 different

groups. The complexity and unique development of the structures of the phallo-

some and proctiger of the male genitalia in each species coupled with lack of

ecological differentiation and with sympatric occurrence suggest that in these

forms the effective mechanism of species Separation is a mechanical barrier

in mating. This interpretation is supported by the fact the females of all spe-

cieS possess a complex bursa copulatrix with an apparently unique development

of internal sclerites in every species. It would be interesting to study these in

detail to see if there is a''lock and key" correspondence of the female and male

genitalia in every Species.

The analyses presented in the preceding paragraphs suggest the following

pattern of evolution and speciation of Paradixa in New Zealand. First, the

problem of whether 1 or 2 lineages entered New Zealand cannot be resolved

definitely at this time because I have not seen extralimital material to deter-

Belkin: Culicidae of New Zealand 33

mine the possible presence of the harrisi and neozelandica groups elsewhere in

the World. However, Tonnoir's illustrations and descriptions suggest that none

of the described Tasmanian and Southeast Australian species belongs to either

group. Therefore, for the present, I am postulating that only 1 lineage of

Paradixa reached and survived in New Zealand. Since tonnoiri possesses the

largest number of generalized features in the male genitalia and in the larva I

believe that this species is closest to the original stock. The first derivative

was harrisi and it was formed probably in geographical isolation somewhere in

the area of the present-day North Island very soon after the arrival of Paradixa

in New Zealand. Further speciation was apparently restricted to the neozeland-

ica group and probably occurred much later. Neozelandica and fuscinervis

may have arisencontemporaneously as aresult of a tripartite geographical iso-

lation of the tonnoiri stock and later become sympatric when the isolated areas

were reunited. However, fuscinervis may have arisen at a later date than

neozelandica either from the latter or from tonnoiri; the dominance of fusciner-

vis might be interpreted as evidence in support of more recent origin of this

species. There is no way to localize the place of origin of either fuscinervis

or neozelandica since they have nearly identical distributions and since relative

abundance and dominance would lead to ambiguous conclusions.

As indicated under Nothodixa, I believe that the absence of Paradixa from

Patagonia and South Chile where Nothodixa is strongly represented is probably

due to a later dispersal of Paradixa, at a time when no Suitable land bridges

existed between New Zealand-Australia and southern South America.

BIONOMICS. In New Zealand, Paradixa is much more common than No-

thodixa probably because the range of its breeding sites is muchwider. Almost

all types of fresh ground waters are utilized except large rivers and the swiftest

small streams. Two of the species, neozelandica and fuscinervis, are conspic-

uous elements of the culicid fauna of New Zealand. However, even these widely

distributed species usually have very small populations.

Paradixa larvae can usually be distinguished from Nothodixa in the field

by the lack of abdominal rosettes projecting above the water surface film. In

natural habitats they normally rest on the edges of the breeding sites with the

body bent in a U, only the head and tail ends in the water. The larvae are

capable of crawling out of water and can move rapidly on the surface film by a

backward looping motion. Pupae can swim but are most frequently found par-

tially or completely out of water but, of course, surrounded by a water film.

Emergence of the adult takes place from the pupa completely out of the water.

Both sexes can be found resting on vegetation in shaded protected places around

the breeding sites. In the laboratory they live only a few days. Males of some

species fly in small swarms, usually near sunset. Eggs are laid on the edge

of the water in a gelatinous mass in which the first instar larvae remain up to

24 hours (Forsyth 1968). The aquatic cycle is completed in about 4 weeks in

the laboratory.

Keys to Species

Adults

a All veins in distal part of wing, particularly ~ 5, with distinct broad

marginal infuscations. . . va we Olle Paseinervis

Veins in distal part of wing at most with very narrow indistinct margin-

Ay OCR Oe gg eer Cane ee re ROU Te Eg gg

Zl)

3(2).

21).

3(2).

Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

Wing without blotch on r-m and base of branches of radial sector .

ee 8. tonnoiri

Wing co more or less distinct blotch on r-m ‘and pase of branches of

badial sGetor 6 ee ee Pe ee 8

Wing blotch small, invading only apex of cell R, otherwise confined to

VEIN mareins . .. . . 6. neozelandica

Wing blotch large, oe? ek of cell R, pase f cells R5 and Rg and

projecting into Cell Ry. wo. a ; ey OS DAPPISi

Male Genitalia

Sidepiece with a winglike claspette lobe ihe from the mesosternal

mMarein . . be ee ee oe aS

Sidepiece ot aie elgcuctte bbe . Be oe ae uk eo So ee

. Sidepiece with a prominent elongate basal tergomesal lobe bearing a

heavy apical spiniform .. . ee . . 6. neozelandica

Sidepiece without prominent elongate basal tergomesal lobe henge 3

. Subapical lobe of sidepiece with at least 1 dark spiniform; clasper ah

only a basal lobe bearing several dark spiniforms . . .7. fuscinervis

Subapical lobe of sidepiece without any dark spiniforms; clasper with

both a basal and a median lobe each bearing several dark spiniforms

. 8. tonnoiri

Pupae

Pinna of trumpet widely flared phn paddle hair 1-P at level of base

of genital lobe .. . : ~ 9. harrisi

Pinna of trumpet narrow wnibaliye not Ttaned: paddle hair 1-P at level

poker middle of genital lobe .

: : ee 6. Heomelantion: 7. fisbinervis: 8. =e

Larvae

Ventrocaudal border of pecten plate with 3-5 very large dark teeth fused

at base; pecten hair 1-S irregularly dendritic; anal segment short,

subequal to length of posterolateral lobe of spiracular apparatus.

‘ : 9. harrisi

Ventrocaudal oe: Aa pecten plate with separate teeth subequal to those

elsewhere; pecten hair 1-S forked or 2, 3-branched; anal segment

long, at least 1.25 length of posterolateral lobe of spiracular appa-

S/S AE tn ek SAIN Cee rm rs See aaa ee a ee eee

Dorsum of thorax without any spicules visible at 100X; antenna much

darker than head capsule .. . se 4 he PUSCINETVIS

Dorsum of thorax with distinct spicules visible at 100X: antenna concol-

Orous With Or lighter (han heaa Capsule 2. ke ee ek ae eo

Basal postpecten spines very large and dark; no differentiated apical

postpecten spicule .... . . 6. neozelandica

Basal postpecten spines very ae ‘light in Hi colon: a | Sonepicsbus dif-

-ferentiated apical postpecten spicule PL Se So ee Gy CONNOITT

Belkin: Culicidae of New Zealand oo

6. Paradixa neozelandica (Tonnoir)

Figs. 1, 8,9, 27

1924. Dixa neozelandica Tonnoir, 1924b:228-229. TYPE: *Holotype 0, Aniseed

Valley, Nelson, N. Z., from small rivulet, 21 Mar 1922, A.L. Tonnoir

[NELS].

Paradixa neozelandica of Belkin (1962:106-107).

Dixa (Paradixa) neozelandica of Tonnoir (1925:311); Edwards (1932:14); Taylor

(1934:7); Nowell (1951:238).

Dixa neozelandica of Cooper and Rapp (1944:251); Miller (1950:45).

ADULTS (figs.1,8; Tonnoir 1924b:fig.2; Belkin 1962:figs. 32,34). Wing:

3.0-4.25 mm. Forefemur: 1.4-1.75 mm. Abdomen: about 1.75-2.25 mm.

As figured here and as described by Belkin (1962:106-107) except for the

following corrections and diagnostic features. General coloration brown with a

variable amount of lighter, yellowish markings. Wing with an inconspicuous

dilute cloud on r-m restricted to vein margins and apex of cell R but not in-

vading cell Rj; no distinct marginal infuscations on other veins, although some

of them, particularly Rs and base of Cu appear darker. Head capsule, including

entire clypeus, dark brown; flagellar segment 1 about 12 (11-13) times as long

as wide and about 1.7 of length of segment 2. Mesonotum light brown on disc

and prescutellar space, dark brown in front and sides, margined with lighter

yellow color on humeral angle and above pleuron; pleuron dark in the center and

below, lighter (yellowish) on upper part along mesonotum and wing base and

along a band from forecoxa to metapleuron; thoracic bristles strong, dark.

Legs brown to light brown, apices of femora and tibiae darkened; distal part of

forecoxa and all of midcoxa and hindcoxa whitish. Haltere largely light, some-

what darkened in middle part of stem.

MALE GENITALIA (fig.8; Tonnoir 1924b:fig.7; Belkin 1962:fig. 33). As

figured here and as described by Belkin (1962:107), with the following cor-

rections and diagnostic features. Sidepiece: A prominent slender short apical

lobe bearing distally 2-4 short heavy spiniforms and several bristles, some

thickened; a prominent thumblike basal mesal lobe bearing apically a long,

very heavy dark spiniform. Clasper with poorly indicated submedian lobe

bearing numerous spiniforms, latter present also distally toward apex of clasp-

er. Phallosome: Prosophallic sclerite very broad, hollowed out in the center

on median projecting lobe. Aedeagal sclerites narrow near base, elongate on

distal part which is bent at an angleof about 90° cephalad. Proximal division of

basal piece very broad. Opisthophallus with a prominent spiculose lateral scle-

rite bearing a thumblike- process at its inner proximal angle and a slender

fingerlike straight process with an apical spiniform from the middle of its

mesal margin. Proctiger: Sclerotizations restricted; a heavily sclerotized

small basal ventrolateral plate produced on its caudomesal angle’into a very

long slender process which bears an apical spiniform and is recurved at base

so that it projects behind proctiger's apex; a poorly sclerotized dorsolateral

plate bearing spicules ventrally and produced at its dorsolateral angle into a

long slender spinelike process strongly recurved cephalad distally and bearing

an apical spiniform and 2-5 subapical cercal bristles.

PUPA (fig.9). Abdomen: 3.1mm. Trumpet: 0.21 mm. Paddle: 0.73 mm.

General morphology and chaetotaxy as figured; diagnostic characters as in the

key; indistinguishable from fuscinervis and tonnoiri. Cephalothorax uniformly

36 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

moderately to lightly pigmented throughout; dorsal surface, except on midline,

with small sparse tubercles; hairs moderately pigmented. Trumpet more or

less barrel-shaped, not widened apically, variable in length; uniformly moder-

ately pigmented, a little brighter than cephalothorax. Abdomen uniformly

lightly to moderately pigmented; tubercles indistinct except laterally; hairs

moderately pigmented. Paddle moderately pigmented; serrations of internal

and external margins variable; hair 1-P somewhat variable in position but al-

ways distad of level of middle of genital lobe. ,

LARVA (fig.9; Tonnoir 1924b:figs.12A,12D). Head: 0.48 mm. Postero-

lateral Lobe: 0.35 mm. Anal Segment: 0.50 mm. General morphology and

chaetotaxy as figured; diagnostic characters as in the key; very similar to

fuscinervis and tonnoiri. Head: Uniformly moderately pigmented a clear

brownish yellow except for darkened antennal tubercle and patches of darkened

areas caudomesad, and black collar; hair pigmentation moderate; hair 11-C

usually with 2,3 forks. Antenna uniformly moderately pigmented, concolorous

with general surface of head capsule. Thorax: Dorsum of posterior part of

prothorax and all of meso- and metathorax with short thickened spicules clearly

visible at 100X and shorter than distance between their bases; sides of all seg-

ments with spicules longer, finer and densely packed; venter of prothorax with

very fine long spicules very densely packed except for posterior part where

they are thickened and more widely spaced; venter of meso- and metathorax

with very fine short spicules densely packed in short transverse lines except

on anterior part of mesothorax where they are similar to those on posterior

part of prothorax; hairs heavily to moderately pigmented, all single. Abdomen:

Spiculation of segment I as on metathorax; spiculationof segments II-VI largely

restricted to midventral areas, spicules progressively longer and more numer-

ous on distal segments, particularly proximad of ambulacral combs on each

segment, dorsal intersegmental areas with shorter heavier spicules; segments

VII, VIII with sparse spicules evident on the sides and dorsum in addition to

the usual long dense ventral spiculation as on preceding segments. Prolegs I, IT

each with about 15-19 large dark hooks in anterior row. Ambulacral combs

with about 7 large spines on each side of median sclerite on segments V, VI

and about 5, 6 spines on segment VII. Hairs strongly to moderately pigmented.

Spiracular Apparatus: Hair 13-S about as long as posterolateral lobe, rather

thin and attenuate apically. Pecten plate moderately to lightly pigmented, with

sparse hairlike spicules in basal part; posterior margin with a row of large

teeth mostly with 2 subequal spines; proximad of these teeth 2 rows of broad

apically fringed spicules; 2,3 very large black basal postpecten spines arising

from a large basal sclerite; apical postpecten spicule not developed; pecten

hair 1-S 2-branched or 2-forked. Anal Segment: Unsclerotized part of segments

IX, X with dense long spiculationon venter as on preceding segments; scleroti-

zation uniformly moderately pigmented except for darkened base of saddle and

apex of process; a few light hairlike spicules on basal saddle and longer similar

spicules on entire surface of caudal process; length of entire sclerotized part

about 1.4-1.5 of posterolateral lobe; postanal spicules ree e ly short, less

than twice the length of cylindrical process.

SYSTEMATICS. Adults of neozelandica are usually readily distinguished

from all other New Zealand dixids by the characteristic infuscation around r-m

of the wing. P. harrisi has a somewhat similar infuscation but it is larger and

more intense than in neozelandica. P. tonnoiri apparently lacks this infuscation

but as noted under this species this may be due to the teneral condition of the 4

known specimens. The degree of infuscation in neozelandica varies consider-

Belkin: Culicidae of New Zealand oF

ably; in addition to the weak cloud on r-m there are more or less distinct in-

fuscations on the distal part of nearly all the veins and at the base of Cu, but

these infuscations rarely approach the condition in fuscinervis and there is no

difficulty usually in separating the 2 species because of the much larger and

denser cloud on r-m in fuscinervis. The variation in wing infuscation in neo-

zelandica does not appear to be correlated with differences in ecological or

geographical origin of the samples. In the series of neozelandica from the colo-

ny established by Forsyth (1968) a number of striking anomalies in the wing ve-

nation have been noted, primarily unilateral, as indicated under Neodixa minu-

ta. I have not seen any such anomalies in adults reared from larvae collected

in nature. As in the case of other species of New Zealand Paradixa, the male

genitalia of neozelandica are very characteristic. There is no significant vari-

ation in the unique development of the phallosome and proctiger complexes. On

the other hand, the lobes and spiniforms of the sidepiece and the spiniforms of

the clasper show a great deal of variation within a single population but as in

the case of wing characters there is no indication of ecological or geographical

differentiation in these structures. P. neozelandica differs from all other New

Zealand paradixines in the development on the sidepiece of a prominent basal

lobe with a heavy apical spiniform. The main body of the lateral sclerite of the

opisthophallus of neozelandica is very strongly developed as a prominent spicu-

late lobe extending laterad. This lobe is homologous with differently developed

structures infuscinervis and tonnoiri but is completely different from the clasp-

ettelike lobe developed from the sidepiece in harrisi.

The larva of neozelandica is very similar to fuscinervis and particularly

tonnoirifrom which it is differentiated primarily by the presence of very strong-

ly developed and very dark postpecten spines arising from a distinct basal scle-

rite. P. fuscinervis has similar but muchsmaller and lighter postpecten spines

and in addition has dark antennae and lacks distinct thoracic spiculation. There

is considerable variation in the development of the pecten teeth and basal post-

pecten spines in neozelandica but no significant overlap in the latter character

with the other 2 species and no indication of anything but individual variation

for both characters. |

P. neozelandica is undoubtedly closely related to fuscinervis and tonnoiri

as indicated by basic similarity in all stages, all 3 species being strikingly dif-

ferentiated only in the male genitalia. In extent of geographical and ecological

range and in abundance and dominance, neozelandica approaches fuscinervis.

This may indicate that it was the second derivative of this lineage, the neoze-

landica complex.

BIONOMICS. P. neozelandica has nearly as wide a range of natural

breeding sites as fuscinervis but appears to be less common. It has been col-

lected in pools, ponds and swamps more frequently than fuscinervis and this may

indicate that it has a preference for quieter water. Several collections are

known from streams with moderate to fast general current but it is possible

that in such situations neozelandica breeds in the quieter parts of the streams.

The light conditions in the breeding sites ranged from deep shade to full sunlight.

In all but 8 of the total of 50 larval collections, neozelandica was associated

with fuscinervis. P. neozelandica was found in 3 of the 4 larval collections of

tonnoiri, in 6 of 10 collections of harrisi, in 7 of 17 collections of Nothodixa

campbelli, but never with Nothodixa septentrionalis.

P. neozelandica apparently does not penetrate into mountainous areas as

much as fuscinervis, but like fuscinervis it has taken advantage of man-made

habitats. It has been collected in ditches and has been found in experimental

38 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

ponds at the Mangere Sewage Treatment Plant in Auckland. As in the case of

fuscinervis, the larval populations of neozelandica are usually small and only a

few adults are found resting in the vegetation surrounding the breeding sites.

P. neozelandica is the only dixine colonized to date (Forsyth 1968). In

this colony, maintained for 5 generations, females oviposited within 72 hours

of emergence and without taking any food. From 25 to 78 eggs were enclosed

in a single circular gelatinous mass which was attached to the walls of the tank

at the water level. The larvae hatched after 4 days but remained in the gelati-

nous matrix up to 24 hours before swimming out. The aquatic cycle lasted an

average of 30 days at 20°C and the adults lived on the average only 4 days.

DISTRIBUTION (fig. 27). Material examined: 1324 specimens; 101°", 1669,

3 adults, 902 larvae, 152 pupae; 131 individual rearings (82 larval, 43 pupal,

6 incomplete).

NEW ZEALAND. North Auckland: Mangamuka Gorge, 2 mi N of Soda

Spring, 21 Dec 1963, BS (NZ 41), 1 lpo (41-111), 2 1p? (41-107, 110), 5 L

[UCLA]; same data (NZ 42), 1 o [UCLA]. Dome Valley, 2 Jan 1965, F (NZ

198B), 10°, 22|[UCLA]. Mangawhai Heads, elev. 12 m, 2 Jan 1965, F (NZ

198A), 12, 3 L[UCLA]. Mahurangi Heads, Martins Bay, elev. 3m, 26 Dec

1964, F (NZ 196), 2 1p (196-102, 103), 1 lp? (196-101), 1 lp (196-104), 19

[UCLA]; 29 Dec 1964, F (NZ 197), 2 lp? (197-101, 103), 1 p? (197-102), 1 L

[UCLA]. Albany, Rosedale Road, elev. 100m, 18 May 1964, F (NZ 182), 1 1p@

(182-123), 1 lp (182-121), 4 L[UCLA]. Bethells Road Swamp, 10 Jan 1965, F

(NZ 200B), 1 po (200B-101), 1 p? (200B-102), 10°, 29, 2 L[UCLA]; 12 Jan

1965, F (NZ 202), 1c, 12, 1 L[UCLA]. Waitakere Ranges, edge of Cascade

Park, 18 Dec 1963, BS (NZ 35), 4 L[UCLA]; elev. 20 m, 15 Nov 1964, F (NZ

195B), 2 L[UCLA]. Nihotupu, 23 Feb 1923, A.L. Tonnoir, 1 A[NELS]. Titi-

rangi Beach, 30 Dec 1963, BS (NZ 64), 6 1p? (64-102, 103, 105-108), 10, 2 9,

17 L, 5 P[UCLA]; 11 Jan 1964 (NZ 94), 1 1p? (94-105), 1 po (94-104), 1 p?

(94-102), 86 L[UCLA]. Titirangi, French Beach, elev. 5m, 4 July 1964, F

(NZ 192), 1c, 2 L[UCLA]. Mangere Sewage Treatment Plant, 9 Mar 1965, F

(NZ 209A, B,C), 1 lpo& (209B-102), 17 1p? (209B-101, 103-111, 114, 115, 123,

126-129), 4 po (209B-113, 116,119,120), 7 p? (209B-112, 117, 118, 121, 122, 124, |

125); 8 0, 38 2 (200A), 30 Ly 1 P.(2098),30 &}) 889, 10 b,.5 P (209C) | UCLA;

Mangere Purification Plant, experimental pond A, 15 June 1965, F (NZ 215),

1 lp? (215-11), 1 po (215-10) [UCLA]; same data except experimental pond B

(NZ 216A), 1 1p? (216A-110), 5 po (216A-100-102, 104, 109), 5 p? (216A-103,

105-108), 1c, 19[UCLA]. Avondale, elev. 60 m, 23 Nov 1964, F (NZ 216B),

1 lp? (216B-102), 1 p? (216B-101), 15¢°, 59 [UCLA]. Tuakau, elev. ca 10 m,

9 June 1964, F (NZ 190), 2 L[UCLA].

Little Barrier Island: Waipawa stream, 75-100 ft from mouth, 6 Mar 1964,

S (NZ 130), 1 1p (130-101), 5 lp? (130-102-104, 107, 108), 1 po (130-105),

1 p@ (130-106), 2c, 29, 40 L, 3 P[UCLA]. Hut Bay Creek, near sea level,

8 Mar 1964, S (NZ 134), 1 L [UCLA].

Great Barrier Island: Port Fitzroy, 12 Mar 1964, S(NZ 138), 42 L[UCLA].

South Auckland: Coromandel, 25 Apr 1964, N (NZ 221), 25 L[UCLA]. Te

Aroha, 10 mi S; 5 June 1965, F (NZ 211), 2 1p? (211-100, 102), 1 po (211-103),

19, 14 L[UCLA].

Gisborne: Ormond, Waihirere Domain, 25-26 Dec 1963, BS (NZ 48), 2 1p

(48-109, 114), 1 p? (48-103), 34 L[UCLA]; 28 Mar 1964, S (NZ 140), 1 Ip

(140-102), 2 po (140-101, 104), 1 1p (140-105), 10 L[UCLA]. Hexton, bridge

H20, S16-41, 26 Dec 1963, BS (NZ 50), 1 1lpo (50-112), 5 lp? (50-113-117),

12 L [UCLA].

Belkin: Culicidae of New Zealand . 39

Wellington: Manunui turnoff between Piriaka and Taumarunui, 7 June 1965,

F (NZ:213), Lipo’ (213104), 2:1p¢ (213-101, 103), 1: p?(213-105), 1 LLVCLA,

Piriaka, elev. 200 m, 5 Jan 1965, F (NZ 199), 1 1p? (199-102), 2 L [UCLA];

28 Mar 1965, F (NZ 210), 2c [UCLA]. Pokaka, 10 Jan 1964, BFS (NZ 86),

8 L [UCLA]. Ohakune, Mountain Road, 9 Jan 1964, BFS (NZ 83), 11 L[UCLA];

pond on bend of Lake Road, 9-10 Jan 1964, BFS (NZ 85B), 4 lpo& (85B-108,

110, 113, 120), 10 1p? (85B-102, 104-107, 109, 115-118), 2 po’ (85B-112, 114),

3:p9 (85B-101, 103, 111), 1 1p (85B-119),-10; 179: L, 3 P [UCLA]; Rangataua

Reserve Lake, elev. 600 m, 6 June 1965, F (NZ 212A), 12 [UCLA]; Ohakune,

8 Mar 1923, A.L. Tonnoir, 1% [BMNH], Mar 1923, T.R. Harris, 1 “|[BMNH],

May-July 1923, T.R. Harris, 19 [BMNH], Feb1924, T.R. Harris, 1 9/BMNH].

Nelson: The Glen, 12 Mar 1964, S (NZ 168), 1 1p? (168-101), 8 L[UCLA].

Wakapuaka, Glen Road, 12 Mar 1964, S (NZ 167), 2 L[UCLA]. Maitai River,

near Pole Ford Bridge, 14 Mar 1964, S (NZ 170), 9 L [UCLA]; at Smith's ford,

14 Mar 1964, S(NZ 171), 9 L[UCLA]. Nelson Reservoir, 12 Mar 1964, S(NZ

162), 7 L [UCLA]. Nelson, near DSIR, 3 Jan 1964, B(NZ 73A & B), 40%, 4L

[UCLA]. Nelson, 3 Dec 1921, A.L. Tonnoir, 2 & paratypes (I: 293, I- 294)

[CANT]; 30 Oct 1923, A.L. Tonnoir, 1 A paratype [NELS]. Annesbrook, elev.

ca 20 m, 10 Mar 1964, S(NZ 158), 1 1p@ (158-101), 7 L[UCLA]. Blick's Val-

ley, 12 Mar 1964, S (NZ 164), 1 po (164-101), 9 L [UCLA]. Dodson's Valley,

elev. ca 68m, 12 Mar 1964, S (NZ 165), 3 L[UCLA]. Todd's Valley, 12 Mar

1964, S (NZ 166), 1 lp? (166-102), 6 L[UCLA]. Aniseed Valley, 21 Mar 1922,

A.L. Tonnoir, & holotype, 1, 2 2 paratypes [NELS]; 5 Jan 1964, BG (NZ

80), 3 lp? (80-106-108), 19 L[UCLA]. Howard Junction, 13 Mar 1964, S (NZ

169), 17 L[UCLA]. Gowanbridge, 9 Feb 1964, S (NZ 122), 5 L [UCLA]. Gow-

an River, road to Lake Rotoroa, 9 Feb 1964, S (NZ 121), 2 p? (121-101, 102),

90 L [UCLA]. New Creek Road, between Murchison and Kawatiri Junction, 8

Feb 1964, S (NZ 116), 1 L [UCLA].

Westland: Hercules Mt, S of Harihari, 7 Feb 1964, S (NZ 115), 1 1p (115-

109), 1 L[UCLA]. Whataroa, 7 Feb 1964, S (NZ 114), 8 L[UCLA].

Canterbury: Christchurch, 28 Nov 1924, 12[USNM]. Purau Creek, 20

Feb 1922, A.L. Tonnoir, 12 paratype [NELS], 19 paratype [CANT]. Timaru,

Scenic Reserve, 24 Jan 1964, S (NZ 100), 3 1p? (100-103, 104,114), 1 1p (100-

110)0 8 GH IUCRAT

Otago: Waianakarua River near bridge of main road, 25 Jan 1964, S (NZ

102), 1 1p (102-120), 1 1p (102-122), 66 L[UCLA]. Dunedin, Blackhead, 23

Feb 1964, N (NZ 217), 1%, 29, 7 L[UCLA]. Queenstown, Ben Lomond, 2

Feb 1964, S(NZ 111), 1 L [UCLA]. |

New Zealand, location uncertain: Horseshoe Lake, 11 Nov 1924, 19 [USNM].

7. Paradixa fuscinervis (Tonnoir)

Figs. 7, 10, 28

1924. Dixa fuscinervis Tonnoir, 1924b:229-230. TYPE: *Holotype “, Nelson,

N.Z., from small rivulet, 5 Sept 1922, A.L. Tonnoir [NELS].

Paradixa fuscinervis of Belkin (1962:109-110).

Dixa (Paradixa) fuscinervis of Edwards (1932:14); Taylor (1934:7); Nowell

(1951:234).

Dixa fuscinervis of Cooper and Rapp (1944:249); Miller (1950:45).

ADULTS (Tonnoir 1924b:fig.4; Belkin 1962:fig.34). Wing: 3.5-4.7 mm.

40 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

Forefemur: 1.7-1.9 mm. Abdomen: about 2.2-2.5 mm. As described by

Belkin (1962:109); extremely similar to neozelandica and differing almost en-

tirely in wing ornamentation. Distal part of all veins from R to Cug with

rather broad marginal infuscations, particularly strong and wide on Cus and

M344; all of Cu, base of 1A and hind margin of wing with more or less broad

marginal infuscations; a large conspicuous dark blotchon r-m extending across

furcation of Rs into cell R and almost touching vein Ry. Somewhat larger than

neozelandica and usually lighter in coloration; clypeus usually lighter apically;

length and proportions of flagellar segments similar; mesonotum light brownon

entire disc except for darker restricted supraalar stripes (no medianor anterior

darkening); lateral mesonotal and pleural markings essentially similar to neo-

zelandica; thoracic bristles weaker and lighter. Legs lighter in color, coxae

Similar to neozelandica.

MALE GENITALIA (fig. 7; Tonnoir 1924b:fig.6; Belkin 1962:fig.33). As

figured here and as described by Belkin (1962) with the following corrections

and diagnostic features. Sidepiece: A short apical lobe bearing 1 or more

Slender spiniforms and several bristles apically; no basal mesal lobe; no basal

tergal bristles. Clasper with a conspicuous submedian lobe bearing about 10

short blunt spiniforms distally; shaft distad of lobe without spiniforms. Phallo-

some: Prosophallic sclerite narrower at base than in neozelandica, with avery

prominent long outstanding median process hollowed out on its posterior face.

Aedeagal sclerite essentially as in neozelandica but more heavily sclerotized

and broader throughout. Proximal division of basal piece less strongly devel-

oped. Lateral sclerite of opisthophallus narrower, rugose laterally but not

Spiculose; its proximal process shorter, broader; its distal process shorter,

stouter, curved and arising farther distad. Proctiger: With strong basal ven-

trolateral sclerotization from which arises first a very long slender curved

spinelike process bearing an apical spiniform, more distally a shorter similar

spinelike process, and cephalad of the latter and at about same level a short

sclerotized thumblike truncate process bearing 2 widely separated apical cer-

cal bristles.

PUPA (fig.10). Abdomen: 3.4 mm. Trumpet: 0.25 mm. Paddle: 0.79 mm.

General morphology and chaetotaxy as figured; diagnostic characters as in

the key; indistinguishable from neozelandica and tonnoiri. Cephalothorax

uniformly moderately pigmented; tubercles apparently larger and more numer-

ous than in neozelandica; hairs lightly pigmented, difficult to see. Trumpet

more or less barrel-shaped, not widened apically, variable in length; uniformly

moderately pigmented, somewhat brighter than cephalothorax. Abdomen mod-

erately to lightly pigmented, sometimes slightly darker laterally; tubercles in-

distinct except laterally; hairs lightly pigmented. Paddle uniformly moderately

pigmented; serrations of internal and external margins variable; hair 1-P dis-

tad of level of middle of genital lobe.

LARVA (fig. 10; Tonnoir 1924b:fig.12C). Head: 0.53 mm. Posterolateral

Lobe: 0.33 mm. Anal Segment: 0.43 mm. General morphology and chaeto-

taxy as figured; diagnostic characters as in the key; very similar to neoze-

landica and tonnoiri, differing from the former primarily-in the following fea-

tures. Head capsule lighter in color. Antenna moderately pigmented, concol-

orous with basal tubercle and markedly darker than general coloration of head

capsule. Thoracic and abdominal spiculation poorly developed, indistinct ex-

cept on venter of thorax and distal abdominal segments; dorsum of thorax with-

out visible spiculation at 100X. Hair13-S somewhat shorter than posterolateral

lobe of Spiracular apparatus. Pecten plate lightly pigmented, poorly differenti-

Belkin: Culicidae of New Zealand 41

ated from unsclerotized part of segment, without distinct basal spicules; mar-

ginal pecten teeth weaker and shorter; basal postpecten spines developed but

weaker, straighter and lighter in color, usually 1 or 2. Anal sclerotization

much more lightly pigmented but darkened in the middle, length about 1.3 of

posterolateral lobe; basal saddle without distinct spicules; caudal process

Shorter, spicules restricted to sides; postanal spicules about twice as long as

cylindrical process.

SYSTEMATICS. Adults of fuscinervis are usually readily distinguished

from all other New Zealand dixids by the characteristic infuscation of the wing.

Nothodixa campbelli and N. philpotti have somewhat similar patterns of wing

infuscation but are easily separated by striking differences in wing spotting

and also by the short antennae. As indicated under neozelandica, that spe-

cies has more or less distinct narrow marginal vein infuscations in addition

to a Small cloud on r-m but in spite of considerable variation in wing infuscation

in fuscinervis there is no significant overlap between the 2 species. The most

reliable character to separate them is the much larger and denser cloud on r-m

in fuscinervis. The variation in wing infuscation in fuscinervis is strictly in-

dividual and does not correlate with any differences in ecology or distribution.

I have not noted any striking anomalies in the wing venation of fuscinervis ina

relatively large series. The male genitalia of fuscinervis have the greatest de-

velopment of the phallosome and proctiger complex of any New Zealand Para-

dixa but basically are of the type found in neozelandica and tonnoiri. There is

no significant variation in the unique development of any of these structures

and particularly in the very prominent median process of the prosophallus, the

very large aedeagal sclerite, and the very broad rugose lateral sclerite of the

opisthophallus and its spines. On the other hand, as in neozelandica, there is

a great deal of variation in the development of the apical lobe of the sidepiece

and the submedian lobe and spiniforms of the clasper. These variations are al-

so strictly individual and do not correlate with differences in ecology or geo-

graphical origin of the different populations.

The larva of fuscinervis is very similar to neozelandica and tonnoiri but

is readily separated from both by the very dark antennae and the absence of

conspicuous spiculation on the thorax. There is considerable variation in the

pigmentation and development of the pecten teeth and spicules and particularly

in the development of the basal postpecten spines. The latter, however, are

always much smaller and lighter in color than in neozelandica and much larger

than in tonnoiri and there is no significant overlap with either species. This

variation of the pecten structures is, as in the other species of the complex,

strictly individual.

As indicated under neozelandica, P. fuscinervis is closely related to this

Species and to tonnoiri. It is the dominant species of the neozelandica complex

and the dominant dixid of New Zealand. Compared to the other widespread spe-

cies (neozelandica), it has a somewhat wider geographical distribution, a dis-

tinctly wider range of breeding sites, and usually predominates in numbers in

mixed populations. This suggests that it is the latest, most modern derivative

of the neozelandica complex.

BIONOMICS. P. fuscinervis has the widest range of breeding sites of all

New Zealand culicids. It has been found in all types of fresh ground waters, in

streams, Seepages, pools, ponds, lakes and even swamps. Twice it was found

breeding ina fire barrel in association with Culex pervigilans and C. quinque-

fasciatus (NZ 45,66). P. fuscinervis apparently prefers slow-moving or still

water but is also found in streams with moderate or even strong current. It is

42 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

most commonly found in partial shade but has also been collected frequently in

full sunlight and more rarely in deep shade. P. fuscinervis has been found in

association with all the other dixids whose immature stages are known, most

frequently with neozelandica, but in about half of some 80 larval collections of

fuscinervis no other dixid was present. In mixed collections fuscinervis is

usually the dominant species.

P. fuscinervis occurs in the hills and mountains as well as in broad val-

leys, plains and coastal areas. It has adapted well to changes in the environ-

ment brought about by man and breeds incities and agriculturalareas. Although

it is the dominant New Zealand dixid its larval populations areseldom large and

only a few adults are found resting in the shade on vegetation surrounding the

breeding sites.

DISTRIBUTION (fig. 28). Material examined: 2708 specimens; 131°, 1799,

8 adults, 2051 larvae, 339 pupae 221 individual rearings (138 larval, 69 pupal,

14 incomplete),

NEW ZEALAND: North Auckland: Mangamuka Gorge, 2 mi N of Soda

Spring, 21 Dec 1963, BS (NZ 41), 5 1p? (41-103, 104, 106, 108,109), 1 po (41-

101), 1 lp (41-105), 46 L [UCLA]; same data (NZ 42), 2 lpo (42-101, 104),

lp? (41-103, 105), 1 po (41-102), 23 L [UCLA]; S slope at bridge, 18 Feb 1965,

F (NZ 206), 1 lpo (206-131), 5 L[UCLA]. Ruakaka, 20 Dec 1963, BS (NZ 36),

1 L[UCLA]. Wellsford, elev. 100 m, 31 May 1964, D. Forsyth (NZ 188), 3 L

[UCLA]; first bridge S of city, 16 Feb 1965, F (NZ 204), 6 L[UCLA]. Dome

Valley, 2 Jan 1965, F (NZ 198B), 12 [UCLA]: bridge : ai Wellsford, 18 Feb

1965, F (NZ 208), 5 L[UCLA]. Albany, Rosedale Road, elev. 100 m, 18 May

1964, F (NZ sn l:lp?.(182-124), 7. [UCLA]. Bethells Road Scan. 10

Jan 1965, F (NZ 200B), 1 po (200B-103), 1 p? (200B-104), 1h, 19, 2L[UCLA].

12 Jan 1965, F (NZ 202), 10, 2 L[UCLA]. Bethells Road, near quarry, 11

Apr 1964, F (NZ 179), 1 lpo& (179-104). Waitakere Ranges, edge of Cascade

Park, 18 Dec 1963, BS (NZ 35), 7 1p? (35-101, 112-117), 6 po (35-102, 104-

106, 109,111), 4 p? (35-103, 107, 108, 110), 10%, 19, 82 L, 6 P[UCLA]; same

general locality, 28 Dec 1963, BS (NZ 58), 1 po (58-101), 30 L, 1 P [UCLA];

Waitakere stream, 20 Feb 1964, F (NZ 177), 1 co [UCLA]; same locality, date

and collector (NZ 178), 1 po (178-101) [UCLA]; Bethells Road at Cascades

turnoff, elev. 20m, 15 Nov 1964, F (NZ 195A & B), 1° (A), 10 L (B)[UCLA];

West Coast Road, first stream W of Scenic Drive, 8 Jan 1964, BS (NZ 81),

1 lpo& (81-101), 2 1p? (81-102, 103), 14 L [UCLA]; West Coast Road, stream

east of inlet to Nihotupu reservoir, 14 Jan 1964, S (NZ 98), 6 L [UCLA]. Swan-

son, Train Valley stream, elev. 166m, 9 May 1964, F (NZ 183), 3 lp (183-

134,135, 138), 2 lp? (183-136, 137), 12 L [UCLA]. Henderson, Opanuku stream,

elev. 65 m, 26 Apr 1964, F (NZ 181), 1 lp (181-125), 9 1p? (181-122, 124,

126-132), 1 1p (181-123), 20, 23 L, 1 P[UCLA]. Oratia, DSIR Field Station,

22 Dec 1963, BS (NZ 45), 3 L [UCLA]: 30 Dec 1963, S (NZ 66), 2 1p? (66-101,

102), 6 L, 3 P [UCLA]. “Titirangi Beach, creek above dam, 30 Dec 1963, BS

(NZ 64), 2 lp? (64-101, 104), 20%, 1 9, 33 L [UCLA]; 11 Jan 1964, BS (NZ 94),

1 lpo (94-107), 1 lp? (94-101), 1 p? (94-103), 1 1p (94-106), 30°, 116 L,8P

[UCLA]; Mahoe Road, elev. 2m, 4 July 1964, F (NZ 193), 2 1p? (193-121, 122),

9 L [UCLA]; same locality, elev. 5m, 29 Nov 1964, F (NZ 195C), 1c, 19

[UCLA]. Titirangi, French Bay, elev. 50 m, 14 June 1964, F (NZ 189B), 1 L

[UCLA]; elev. 5m, 4 July 1964, F (NZ 192B), 6 L[UCLA]. Avondale, elev.

60 m, 23 Nov 1964, F (NZ 216B), 1 p? (216B-101) [UCLA].

Little Barrier Island: Stream between Te Waikohare and Tirikikawa, elev.

7m, 9 Oct 1963, S(NZ 11), 1 1p? (11-111), 1 1p (11-112) [UCLA]. Waipawa

Belkin: Culicidae of New Zealand 43

stream, 75-100 ft from mouth, 6 Mar 1964, S (NZ 130), 5 L[UCLA]. Tiriki-

kawa stream, elev. 30 m, 8 Mar 1964, S (NZ 131), 2 lpo (131-101, 103);;1: lpe

(131-104), 1 po (131- 102), 16 L [UCLA]. Hut Bay creek, near sea level, 8

Mar 1964, S (NZ 134), I po (134-101), 4 L [UCLA].

Great Barrier Island: Port Fitzroy, 12 Mar 1964, S (NZ 138), 3 po (138-

102, 108,109), 1 p? (138-110), 95 L [UCLA].

South Auckland: Te Aroha, 10 miS, 5 June 1965, F (NZ 211), 3 L[UCLA].

Rotorua, Kuirau Hot Springs Reserve: 23 and 27 Dec 1963, BS (NZ 46), 1 lpo&

(46-140) [UCLA].

Gisborne: Ormond, Waihirere Domain, 25-26 Dec 1963, BS (NZ 48), llpo

(48-104), 5 1p? (48- 101, 106, 107, 110,112), 3 po (48-102, 1 Ut 113), 1 lp (48-

108), 81°L,° bP (UCLA I; 28 Mar 1964, S (NZ 140), 1 p? (140- 103), ie. 8 bk,

4p [UCLA]. Hexton, at bridge H20, S16- 41, 26 Dec 1963, BS (NZ 50), 1 1p?

(50-111), 3,1 P [UCLA]. Urewera National Park, Orangihikoa stream,

elev. 2450 ft, 30 Mar 1964, S (NZ 141), 2 lp? (141- 101, 104), 1 po (141- 102),

4 L[UCLA]; Mokau Falls, 27 Dec 1963, BS (NZ 55), 1 L [UCLA]; Aniwaniwa

Falls, 27 Dec 1963, BS (NZ 52); 8:19 (52- 101-104;106, 107), 3: oo (1 O)- 78 hs,

6 P [UCLA|. Terapatiki, 27 Dec 1963, BS (NZ 51), 1 po (p14 101), 1 L[UCLA|].

Wellington: Mapiu, 7 June 1965, F (NZ 214), 1 L[UCLA]. Between Piri-

aka and Taumarunui, 7 June 1965, F (NZ 213), 1 po (213-102), 10 L [UCLA].

Piriaka, elev. 200 m, 5 Jan 1965, F (NZ 199), 1 1p@ (199- 103), 1 p? (199-

105), 9 i [UCLA]; 28 Mar 1965, F (NZ 210), 1° [UCLA]. Pokaka, 10 Jan 1964,

BFS (NZ 87), 10 1p? (87-101, 108- 113, 125-137); S00 (87-103- 107), 2 1p (87-

102, 144)301 C9, OT L, 14 P [UCLA]; old road junction, 10 Jan 1964, BFS

(NZ 86), 3 1po’ (86-102-105), 1 1p? (86-106), 17 L[UCLA]. Ohakune, Moun-

tain Road, 9 Jan 1964, BFS (NZ 83), 41po (83-106, 109, 110, 115), 10 1p? (83-

103-105, 107, 108, 111-114, 116), 1 po (83-125), 128 L, 18 P[UCLA]; 26 Jan

1964, F (NZ 175), 4 L[UCLA]; Rangataua Reserve Lake, 9 Jan 1964, BFS (NZ

84), 1 L[UCLA]; pond at bend of Lake Road, 9-10 Jan 1964, BFS (NZ 85B),

17 L[UCLA]. Ohakune, Sept 1922, T.R. Harris, 19 [BMNH], 10 Nov 1922,

T.R. Harris, 1? [BMNH], 15 Dec 1922-15 Jan 1923, T.R. Harris, 1 9[BMNH].

Nelson: Riwaka River, North branch, Crystal Pool, 4 Jan 1964, BF (NZ

74), 1 Ipo (74-105), 7 1p9 (74-101-104, 106-108), 40°, 89, 21L, 22 P[UCLA];

Riwaka River valley, north branch, 4 Jan 1964, BG (NZ To) ops, 81 P

[UCLA]. The Glen, 12 Mar 1964, S (NZ 168), 1 lp? (168-102), 4 Do (168- 105,

106, 108,109), 1 De (168-107), 32 L [UCLA]. Wakapuaka, Glen Road, 12 Mar

1964, S (NZ 167), 1 lpo (167-101), 8 L[UCLA]. Todd's Valley, 12 Mar 1964,

S (NZ 166), 1 1lpo (166-101), 59 L [UCLA]. Dodson's Valley, 12 Mar 1964, S

(NZ 165), 34 L[UCLA]. Blick's Valley, 12 Mar 1964, S (NZ 164), 55 L [UCLA].

Nelson Reservoir, 12 Mar 1964, S (NZ 163), 7 L [UCLAI. Nelson, N Branch

Maitai River, elev. ca 400 ft, i1 Feb 1964, S (NZ 123), 1 o [UCLA]; junction

Maitai River and Sclanders stream, 11 Feb 1964, 8(NZ 125), 1 4p? (125-102),

2 po (125-103, 105), 2 p? (125-101, 106), 11p (125- 104), 73 L, 3 P [UCLA];

South Branch Maitai River, 11 Feb 1964, S (NZ 127), 5 L [UCLA]; Maitai

River, near Pole Ford, 14 Mar 1964, S(NZ 170), 24 L[UCLA]; Maitai River

and Dod's Creek, 14 Mar 1964, S (NZ 171), 34 L [UCLA]; Cawthron Park,

elev. ca 450 ft, 11 Feb 1964, S(NZ 124), 36 L[UCLA]; near DSIR, 3 Jan1964,

BG (NZ 73A, B), 1 1lpo (73-108), 6 lp (73-101-104, 107, 109), 2 1p (73-105,

106), 150°, 139, 112 L, 19 P[UCLA]. Nelson, 10 Aug 1922, A.L. Tonnoir,

1 o paratype [NELS], 5 Sept 1922, A.L. Tonnoir, & holotype, 1 A paratype

[NELS], 20 Sept 1923, A.L. Tonnoir, 1 A paratype [NELS], 1 o paratype

(I- 299) [CANT], 28 Sept 1923, A.L. Tonnoir, 1% paratype [BMNH], 7 Dec

AG Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

1921, A.L. Tonnoir, 1 o paratype (I- 301) [CANT]. Ngawhatu Mental Hospital,

11 Mar 1964, S (NZ 159), 1 lpo (159-101), 3 L[UCLA]. Annesbrook, elev. ca

20 m, 10 Mar 1964, S (NZ 158), 6 1p (158-102-104, 106-108), 1 po‘ (158-105),

57 L, 3 P [UCLA]. Brightwater, 11 Mar 1964, S (NZ 161), 1 lpo (161-101),

2 L[UCLA]. Aniseed Valley, 5 Jan 1964, BG (NZ 79), 9 L, 1 P[UCLA]; same

locality and date (NZ 80), 2 1p? (80-101, 105), 20 L[UCLA]. Lake Rotoiti, 9

Feb 1964, S (NZ 117), 1 po (117-108), 12 L [UCLA]. Howard Junction, 1 mi

E, 13 Mar 1964, S (NZ 169), 4 L[UCLA]. Gowanbridge, 9 Feb 1964, S (NZ

122), 12 L [UCLA]. New Creek Road, between Murchison and Kawatiri Junc-

tion, 8 Feb 1964, S (NZ 116), 2 1p? (116-102, 103), 2 pot (116-104, 105), 1p

(116-101), 1 1p (116-107), 36 L[UCLA]. Reefton, Jan, A.L. Tonnoir, A[Ton-

noir 1924b]. Springs Junction, elev. ca 560 m, 9 Mar 1964, S (NZ 156), 9 L

[UCLA ].

Marlborough: Kaikoura, 23 Feb 1922, A.L. Tonnoir, 1 A [NELS].

Westland: Lake Brunner [as Moana], 16-21 Dec 1922, A.L. Tonnoir, 2 9

[CANT]. Hercules Mt vicinity, 7 Feb 1964, S (NZ 115), 1 1p? (115-102), 1 p@

(115-110), 15 L [UCLA]. Whataroa, 7 Feb 1964, S (NZ 114), 1¢ (114-108),

7L[UCLA]. Waiho, A.L. Tonnoir, 17 Jan 1922, 1 o paratype (I- 304) [CANT],

19 Jan 1922, 1A paratype [NELS]; 30 Jan 1922, 4 A paratypes [NELS], 1 o pa-

ratype (I- 298), 1° paratype (I- 302) [CANT]. Bridge Flat-Haast Road, 18 Dec

1966, N (NZ 224), 7 L [UCLA].

Canterbury: Christchurch, 13 July 1924, J.W. Campbell, 1 0% [CANT].

Governors Bay, 3 Jan 1964, BD (NZ 72), 2 lpo& (72-104, 107), 1 1p? (72-106),

2 po (72-101, 108), 31 L[UCLA]. Purau Creek, 20 Feb 1922, A.L. Tonnoir,

1 2 paratype (I- 303) [CANT]; Purau Creek [as Christchurch--R. Purau], 22 Oct

1922, J.W. Campbell, 1 o% [BMNH].

Otago: Ben Lomond, Queenstown, 2 Feb 1964, S (NZ 111), 2 1p? (111-102,

105), 1 po (111-104), 52 L, 2 P[UCLA]. Waianakarua River, 25 Jan 1964, S

(NZ 102), 6 1p (102-107, 109, 116, 118, 119, 124), 2 po’ (102-105, 114), 11 p?

(102-101-104, 106, 108, 110-113, 117), 3 1p (102-115, 121, 123), 53L, 3P[UCLA].

Dunedin, Botanical Gardens, 2 Jan 1964, BN (NZ 96), 1 L [UCLA]. Black Gul-

ly, 31 Jan 1964, S (NZ 109), 1 1p? (109-105), 1 L[UCLA]. Clinton, 9 mi N,

26 Jan 1964, S(NZ 103), 1 1po (103-101), 1 1p (103-103), 11 L [UCLA].

New Zealand, localities uncertain: Horseshoe Lake, 11 Nov 1924, collec-

tor unknown, 1 o [USNM]. Locality and collector not specified, 28 Nov 1924,

1 9 [USNM].

8. Paradixa tonnoiri Belkin, n. sp.

Figs. 7, 11, 29

TYPES: Holotype o (NZ 115-108) with associated pupal skin and genitalia

Slide, vicinity of Hercules Mt., 7-8 miS of Harihari on route 6 (Westland), 7

Feb 1964, D.A. Schroeder [NELS]. Allotype 2 (NZ 115-105) with associated

larval and pupal skins, some data as above [NELS]. Paratypes: same data as

the holotype, 1 o (NZ 115-104) with associated pupal skin and genitalia slide,

1 2 (NZ 115-101) with associated larval and pupal skins, 40 L (NZ 115) [UCLA];

2 L (NZ 115) [USNM]; 2 L (NZ 115) [BMNH]; 2 L (NZ 115) [NELS].

ADULTS. Wing: 2.5-2.7 mm. Forefemur: 1.2-1.3 mm. Abdomen: about

1.7mm. Very similar to neozelandica as described by Belkin (1962:106-107),

differing in the following diagnostic features. Wing without any distinct vein in-

Belkin: Culicidae of New Zealand 45

fuscations or any indication of a cloud on r-m. Mesonotum with light brown

ground color and with a long anterior median stripe and a pair of long supraalar

stripes of dark brown. Legs more uniformly dark. Haltere darkened on knob.

Flagellar segments 1 and 2 as in neozelandica.

MALE GENITALIA (fig. 7). Segment IX: Moderately sclerotized. Tergite

with rather distinct median caudal lobe bearing about 25-30 thin bristles, chief-

ly in distal part; lateral lobe inconspicuous, short and with about 3 thinbristles.

Sternite with about 5,6 heavier bristles on each side. Sidepiece: Short and

broad; uniformly densely spiculate; no basal lobe; a prominent small elongate

apical lobe bearing 5,6 short apical and subapical bristles; dorsal surface with

3,4 short thin bristles near base and 3 long thickened bristles near middle;

lateral surface with 1 very long thickened subapical bristle; ventral surface with

about 12,13 long bristles. Clasper about as long as sidepiece; irregular in

outline; with subbasal and submedian lobes each with 3-5 spiniforms; apex with-

out spiniforms. Phallosome: Prosophallus similar to fuscinervis but its median

process broader and less prominent. Aedeagal sclerite very narrow and capi-

tate distally. Lateral sclerite of opisthophallus much smaller than in neozeland-

ica and fuscinervis but strongly bulging laterally; proximal process not developed;

distal process very large, very broad at base and with thumblike distal narrowed

part with spiniform. Proctiger: Basal ventrolateral sclerotization very strong,

bearing a short basal sternal elongate process with elongate slender apical

spiniform, followed tergally by 2 recurved long spinelike processes each with a

shorter heavier apical spiniform, and a short tergal cercal lobe bearing a slen-

der apical bristle.

PUPA (fig.11). Abdomen: 2.7mm. Trumpet: 0.20 mm. Paddle: 0.65

mm. General morphology and chaetotaxy as figured; diagnostic characters as

in the key; indistinguishable from neozelandica and fuscinervis. Cephalothorax

uniformly moderately to lightly pigmented; tubercles apparently more numerous

than in neozelandica; hairs lightly pigmented. Trumpet more or less barrel-

shaped, not widened apically, quite variable in length; uniformly moderately

pigmented, a little brighter than cephalothorax. Abdomen lightly to moderately

pigmented, sometimes slightly darker laterally; tubercles indistinct except

laterally; hairs lightly pigmented. Paddle uniformly moderately pigmented;

serrations of internal and external margins variable; hair 1-P distad of level of

middle of genital lobe.

LARVA (fig.11). Head: 0.53 mm. Posterolateral Lobe: 0.33 mm. Anal

Segment: 0.51 mm. General morphology and chaetotaxy as figured; diagnostic

characters as in the key; very similar to neozelandica and fuscinervis, differing

from the former chiefly in the following characters. Head capsule moderately

pigmented. Antenna moderately to lightly pigmented, sometimes slightly dark-

ened on apex, concolorous with or lighter than general surface of head capsule.

Thoracic and abdominal spiculation similar to neozelandica except that spic-

ules of thoracic dorsum are usually thinner, longer, lighter and much more

dense, visible at 100X; dorsal intersegmental spicules longer and more hair-

like. Hair 13-S usually distinctly shorter than posterolateral lobe of spiracular

apparatus. Pecten plate lightly to moderately pigmented, distinctly differenti-

ated from unsclerotized integument; hairlike spicules more distinct and usually

present on entire plate; marginal pecten teeth similar to neozelandica but

shorter and sometimes with additional spines, frequently heavily pigmented;

submarginal spicules varied, frequently inconspicuous and in a single row;

usually 1,2 very small light basal postpecten spines; a broad, apically-fringed

large apical postpecten spicule. Anal sclerotization usually lighter in color but

46 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

darkened in the middle, length about 1.5-1.6 of posterolateral lobe; spiculation

of saddle and caudal process more conspicuous; postanal spicules about twice

as long as cylindrical process.

SYSTEMATICS. The adults of tonnoiri are very similar to neozelandica

and could be easily confused with specimens of tne latter with faintly marked

wings. The separation of these 2 species on external color characters of the

adults may bevery tenuous since all 4 known specimens of tonnoiri are teneral

and their color pattern may not be fully developed. The male genitalia of ton-

noiri are very characteristic and unique as in other species of New Zealand

Paradixa, however they clearly belong to the same type as those of neozeland-

ica and fuscinervis and not to the type of harrisi. One similarity is evident

with harrisi in the development of basal tergal hairs suggesting anincipient bas-

al lobe on the sidepiece. The phallosome and proctiger complexes of tonnoiri

are the smallest and simplest of all New Zealand Paradixa. There is a con-

Siderable variation even in the only 2 known specimens in the number and dis-

tribution of spiniforms on the clasper and in the number of bristles in the incip-

ient basal lobe of the sidepiece.

The larvae of tonnoiri resemble neozelandica in the color of the antennae

and in the spiculation of the body but are more like fuscinervis in the speciali-

zations of the pecten. There is considerable variation in the development and

pigmentation of the pecten teeth in the fair-sized sample of larvae of tonnoiri

but no significant overlap with fuscinervis in the diagnostic characters. The 4

larvae from the Bridge Flat population (NZ 224) all have very dark pecten teeth

but in all other features resemble closely the other populations in which dark

pecten teeth are uncommon.

As indicated under neozelandica, P. tonnoiri is closely related to this

species and to fuscinervis. The rather wide known distribution on South Island,

the apparent rarity and the presence of considerable variation within and be-

tween the disjunct populations of tonnoiri are suggestive of a relict population

structure in this species. P. tonnoiri also seems to be the least modified mem-

ber of the neozelandica complex on the basis of the male genitalia and larval

features. It appears, therefore, that tonnoiri represents the earliest develop-

ment of the neozelandica lineage in New Zealand.

BIONOMICS. All 4 collections of immature stages of tonnoiri were made

in partial shade in slow-moving or standing water with abundant vegetation in

broad river valleys. In 3 of these collections tonnoiri was associated with both

neozelandica and fuscinervis and with the latter only in the fourth collection.

In only 1 collection (NZ 115) did it appear to be the dominant species.

DISTRIBUTION (fig. 29). Material examined: 88 specimens; 2, 29, 80

larvae, 4 pupae; 4 individual rearings (2 larval, 2 pupal).

NEW ZEALAND: Nelson: Howard Junction, 1 mi E, along Buller River,

13 Mar 1964, S(NZ 169), 7 L[UCLA]. :

Westland: vicinity of Hercules Mt., 7-8 miS of Harihari on route 6, 7 Feb

1964, S(NZ 115), & holotype, 2 allotype, 2 L paratypes [NELS], 10, 19, 40

L paratypes [UCLA]; 2 L paratypes [USNM]; 2 L paratypes [BMNH]. Bridge

Flat-Haast Rd (NZ 81), 18 Dec 1966, N (NZ 224), 4 L[UCLA].

Otago: Waianakarua River, near bridge of highway 1, 25 Jan 1964, S (NZ

102), 20 L[UCLA].

9. Paradixa harrisi (Tonnoir)

Figs, 7,12, 29

Belkin: Culicidae of New Zealand 47

1925. Dixa harrisi Tonnoir, 1925:311. TYPE: *Holotypeo with genitalia mount

_ (I- 296), Ohakune (Wellington), N. Z., Mar 1923, T.R. Harris [CANT].

Paradixa harrisi of Belkin (1962:108).

Dixa (Paradixa) harrisi of Edwards (1932:14); Taylor (1934:7); Nowell (1951:

280).

Dixa harrisi of Cooper and Rapp (1944:250); Miller (1950:45).

ADULTS (Tonnoir 1925:fig.1; Belkin 1962:figs. 32,34). Wing: 3.0 mm.

Forefemur: 1.25 mm. Abdomen: about 2.0 mm. As described by Belkin

(1962:108); extremely similar to neozelandica and differing from it chiefly in

the darker coloration and wing markings. Wing marked as in neozelandica ex-

cept that cloud on r-mvery intense black instead of diffuse brown and much lar-

ger, invading apex of cell R, base of cells R5 and R3 and projecting into cell Rj.

Head darker but clypeus light in distal half; flagellar segment 1 shorter, about

8,9 times as long as wide and about 1.7 of length of segment 2. Thorax uni-

formly dark except for a lateral light area in upper part of pleuron and wing

base; darker supraalar stripes sometimes evident; bristles weaker and lighter.

Hindfemur whitish; hindtibia lighter in middle.

MALE GENITALIA (fig.7; Tonnoir 1925:fig.1). As figured here and as

described by Belkin (1962:108), with the following corrections and diagnostic

features. Sidepiece: Sternomesal margin produced as a large flat winglike

lobe, the 2 lateral lobes narrowly connected across midline at base; a very

short inconspicuous apical process with 2,3 apical bristles; a barely indicated

basal tergomesal lobe bearing 3 moderately long bristles (as in tonnoiri); 3, 4

long distal tergal bristles. Clasper with a distinct subbasal lobe bearing 3-5

moderately long blunt spiniforms; distal part of shaft with 2,3 similar spini-

forms at least 1 of which is apical. Phallosome: Markedly different from other

New Zealand species. Prosophallus with a very long distal median flat process

crenulate apically, enclosing asmall ovoid unsclerotized area and extending be-

tween the 2 aedeagal sclerites. Aedeagal sclerite very narrow and poorly scle-

rotized. Lateral sclerite of opisthophallus narrow, slightly concave laterally;

its external proximal angle produced as a blunt process; its mesal margin with

a very large recurved long sharp spine (without apical spiniform), followed dis-

tally by a Shorter, straighter, narrower process bearing an apical spiniform.

Proctiger: Basolateral sclerotization strong, its mesal ventral margin pro-

duced into a long, branched process, the basal branch short, both branches

spinelike and bearing each a heavy apical spiniform; dorsolateral sclerotiza-

tion joined to basolateral and nearly meeting its mate middorsally, produced

into a spinelike process with curved apical part bearing a distal heavy spini-

form and 2 subapical cercal bristles.

PUPA (fig.12). Abdomen: 2.8 mm. Trumpet: 0.21 mm. Paddle: 0.71 mm.

General morphology and chaetotaxy as figured; diagnostic characters as in

the key; readily separated from other New Zealand Paradixa. Cephalothorax

uniformly moderately to lightly pigmented; tubercles largely restricted to dor-

sal surface; hairs lightly to moderately pigmented. Trumpet distinctly wid-

ened apically, not barrel-shaped, pinna very wide; uniformly moderately pig-

mented, somewhat brighter than cephalothorax. Abdomen lightly to very

lightly pigmented, somewhat darker laterally; tubercles indistinct except later-

ally; hairs lightly pigmented. Paddle with distal part narrower than in other

species; moderately to lightly pigmented; serrations of internal and external

margins variable; apex frequently with more than 1 heavy tooth; hair 1-P at

level of the base of genital lobe.

48 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

LARVA (fig.12). Head: 0.43 mm. Posterolateral Lobe: 0.28 mm. Anal

Segment: 0.28 mm. General morphology and chaetotaxy as figured; diag-

nostic characters as in the key; in general similar to neozelandica, fuscinervis

and tonnoiri but differing markedly from them in the following features. Head:

Uniformly moderately pigmented with darker areas as in neozelandica; hair

pigmentation moderate; hair 11-C usually with 3,4 forks. Antenna moderately

pigmented, concolorous with or slightly darker than head capsule; basal curva-

ture less distinct; hairlike spicules of ventral surface shorter and heavier.

Thorax with spiculation essentially as in neozelandica; hairs somewhat lighter.

Abdomen: Spiculation pattern essentially as in neozelandica but spicules much

shorter and more numerous, becoming heavier and more widely-spaced on sides

and dorsum of VI-VIII; dorsal intersegmental spicules conspicuous at 100X;

proleg hooks and ambulacral spines as in neozelandica but smaller; hairs some-

what lighter. Spiracular Apparatus: Base (caudal part) of sclerite of antero-

median lobe with short heavy spinelike spicules; median area with separate

large lateraland small median sclerites, lateral sclerites with numerous heavy

spinelike spicules, median sclerite with only a few smaller spicules. Hair 13-S

distinctly longer than posterolateral lobe. Pecten plate moderately pigmented;

with scattered hairlike spicules in basal half; dorsal part of posterior margin

with about 10 darkly pigmented simple teeth of moderate size, ventral part of

posterior margin near apex with 3-7 very large heavy black spinelike teeth

fused at base; proximad of these along posterior margin several Separate

smaller spinelike dark teeth or normal branched pecten teeth; 3 rows of sub-

marginal broad apically-fringed large spicules; no basal postpecten spines or

apical postpecten spicules; pecten hair 1-S with 9-15 dendritic branches. Anal

Segment: Unsclerotized part of segments IX, X densely covered with very long

hairlike spicules; sclerotized part very short, about equal in length to postero-

lateral lobe of spiracular apparatus, heavily pigmented; saddle with a subme-

dian transverse patch of heavy slender spinelike spicules; caudal process with

large number of similar but longer spicules ventrally and 1-3 laterally; post-

anal spicules almost half as long as cylindrical process.

SYSTEMATICS. Superficially, the adults of harrisi are very similar to

neozelandica from which however they can be differentiated easily by the large

saturated dark blotch on r-m, the darker coloration of the thorax and the light

hindfemur. I am recognizing a separate group for harrisi because of striking

characters in the male genitalia (especially the claspettelike lobe of the side-

piece and the prosophallus), the larva (spiracular apparatus, pecten and anal

segment) and the pupa (flared trumpet and position of hair 1 on the paddle).

As in the case of tonnoiri, the rather wide known distribution, in this

case on North Island, the apparent rarity, and the considerable intra- and inter-

population larval variation in harrisi suggest a relict population structure in

this species. This similarity probably indicates that tonnoiri and harrisi were

developed contemporaneously, possibly from the same stock. Itis also possi-

ble that harrisi represents a completely different lineage which arrived in New

Zealand contemporaneously with the neozelandica lineage. As indicated in the .

discussion of the systematics of Paradixa, I favor the first explanation.

BIONOMICS. All 10 collections of immature stages of harrisi came from

the margins or isolated pools of small streams in narrow valleys always in

partial to deep shade. The water was still or only slowly moving. The popu-

lations of harrisi were always very small except in 1 instance on Great Barrier

Island (NZ 138) where this species was the dominant one in association with

neozelandica, fuscinervis and Nothodixa campbelli. It was associated with

Belkin: Culicidae of New Zealand 49

fuscinervis on 4 other occasions and with both fuscinervis and neozelandica in

2 collections in both of which neozelandica was dominant. In 2 collections it was

associated with N. campbelli; in 1 of these neozelandica and fuscinervis were

also present.

Adults of harrisi were not encountered in the field during the present

survey and Tonnoir did not record the circumstances of the capture of the type

series by C. T. Harris.

DISTRIBUTION (fig. 29). Material examined: 155 specimens; 150, 16 9,

94 larvae, 30 pupae; 18 individual rearings (2 larval, 16 pupal).

NEW ZEALAND. North Auckland: Mangamuka Gorge, elev. ca 500 m, 18

Feb 1965, F (NZ 205), 1 L[UCLA]; S slope at bridge, 18 Feb 1965, F (NZ 206),

1 L[UCLA]. Brynderwyn Roadside, S slope, elev. ca 330 m, 18 Feb 1965, F

(NZ 207), 1 p? (207-133), 1 p (207-131), 1 P (207-132) [UCLA]. Titirangi

Beach, creek above dam, 11 Jan 1964, BS (NZ 94), 1%, 3 L[UCLA]. Titiran-

gi, French Beach, elev. 50 m, 14 June 1964, F (NZ 189B), 1 lp (189B-131),

11[UCLA].

Little Barrier Island: Waipawa stream, 75-100 ft from mouth, 6 Mar 1964,

S (NZ 130) 19. Tirikikawa stream, elev. 30 m, 8 Mar 1964, S(NZ 131), 1p?

(131-105). Hut Bay Creek, near sea level, 8 Mar 1964, S(NZ 134), 1 p9? (134-

102), 1p, 7 L [UCLA].

Great Barrier Island: Port Fitzroy, 12 Mar 1964, S (NZ 138), 1 po (138-

101), 2 L[NELS]; 1 po (138-104), 2 L[USNM]; 1 p¢ (138-106), 2 L [BMNH];

2 po (138-103, 114), 1 lp? (138-115), 5 p? (138-107, 111-113, 116), 2 o& (650525-

33,94); 17 Py. 72. (UCLA.

- South Auckland: Te Aroha, 10 miS, 5 June 1965, F (NZ 211), 22, 2p,

1 L [UCLA].

Wellington: Ohakune, March 1923, T.R. Harris, o holotype [CANT], 2,

2° [BMNH]; Nov 1923, T.R. Harris, 1 o [BMNH]; Mountain Road, 9 Jan1964,

BFS (NZ 83), 1 po (83-101), 1 p? (83-102) [UCLA].

SUBFAMILY CULICINAE

The True Mosquitoes

The Culicinae are readily recognized from the Dixinae and Chaoborinae in

all stages by the characters given in the key to the subfamilies.

TERMINOLOGY. The descriptive terminology and abbreviations used by

Belkin (1962:547-561) for all stages is followed exceptfor a few minor changes.

On the head of the adults the interocular space is used here for the median

space between the eyes below the vertex; the frons is nowrestricted to the area

on the front of the head between the eyes and the antennae; interorbital bristles

are substituted for frontal bristles; the orbital bristles are divided into the up-

per orbitals on the vertex and the lower orbitals on the sides. On the wing the

point of branching of vein R into veins Rj and Rs is designated as the furcation

of R, the point of branching of vein Rs into veins R2+3 and R4g,5 as the furca-

tion of Rs, and so on for the furcation of R943, M and Cu. In the male genitalia

the term opisthophallus is introduced; it is explained in the discussion of the

male genitalia of the Dixinae.

SYSTEMATICS AND BIOGEOGRAPHY. The details of the affinities and

systematics of the New Zealand Culicinae are discussed under each tribe be-

low. Only the general features of this fauna are reviewed here. The fauna is

harmonious, relatively extensive, and largely endemic at the species group

50 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

level.

Five of the 10 widespread tribes of the Culicinae recognized by Belkin

(1962:117) are represented in New Zealand. Of the 7 tribes represented in the

relatively well known fauna of Victoria, Australia (Dobrotworsky 1965), only

the Anophelini and Aedeomyiini have not been found in New Zealand.

There are 2 endemic monotypic genera (Opifex in the Aedini and Maori-

goeldia in the Sabethini) and 2 endemic monotypic subgenera (Austromansonia

in Coquillettidia; and Nothoskusea in Aedes). The other subgenera represented

by indigenous species in New Zealand are generally considered to be the prim-

itive ones in their respective genera (the nominate subgenus Culex in the Culi-

cini; Climacura in Culiseta; the nominate subgenus Coquillettidia in the Man-

soniini; Halaedes and Ochlerotatus in Aedes).

Eleven of the 14 species are endemic to New Zealand. The ubiquitous trop-

ical house mosquito, Culex quinquefasciatus, is undoubtedly introduced and the

widespread Aedes (Finlaya) notoscriptus is also considered here to be a recent

human introduction although the evidence supporting this is equivocal. The

third non-endemic species, Aedes (Halaedes) australis, is considered here to

have reached New Zealand by natural means and not human introduction.

Except in the Culicini, each endemic species group is represented in New

Zealand by a single species and without any indication of geographical differen-

tiation. In the pervigilans complex of Culex, 2 ecologically isolated species,

asteliae in leaf axils and rotoruae in thermal waters, are recognized in addi-

tion to the widespread pervigilans which exhibits some geographical differenti-

ation in its peripheral populations. There is a suggestion of hybridization be-

tween pervigilans and the introduced quinquefasciatus.

There is no indication of any affinity between the Culicinae of New Zealand

and of South Chile-Patagonia. In general the affinities are primarily with the

fauna of Southeast Australia-Tasmania; definitely so in case of Opifex, Aedes

(Nothoskusea), Aedes (Halaedes), Culiseta (Climacura) and Coquillettidia (C.),

but not as clearly in case of Coquillettidia (Austromansonia), Aedes (Ochlerota-

tus) and Culex (C.). In most instances the New Zealand representatives appear

to have retained more primitive features than their Australian-Tasmanian rela-

tives and are considered to be the earliest derivatives of their respective phy-

lads.

Avery striking feature of the New Zealand Culicinae is the presence of bris-

tles on the lower surface of the base of vein Sc not only in Culiseta but in Co-

quillettidia (Austromansonia), Opifex and Aedes (Nothoskusea), in the female

only in the latter. This character appears to have been a primitive feature in

the subfamily Culicinae. Equally striking is the presence of spicules on the

outer surface of all the claws of the females (sometimes males also) in at least

one member of each tribe represented in New Zealand. Similar spicules are

found in all New Zealand Dixinae and I consider them to be a primitive feature

in the Culicinae. The other apparent important primitive features retained in

the Culicinae of New Zealand are: (1) in the adults, strong development of the

scape of the antenna in Opifex, wide interorbital space on the head of Opifex,

Aedes (Nothoskusea) and Aedes (Halaedes), and strong development of bristles

and poor development of scales on the head, thorax and abdomen in the same

groups, (2) in the pupa of Opifex the similarity of hair 9-VII to hair 9 on pre-

ceding abdominal segments.

Contrary to the generally accepted view that short male palps and non-plu-

mose antennae are derived characters, I suggest that the primitive state in the

Culicinae is more likely to have been retained in the females and therefore pos-

Belkin: Culicidae of New Zealand 51

tulate the following as the primitive condition in the subfamily: (1) antenna sim-

ilar inthe 2 sexes, with flagellar whorls poorly developed in both, (2) palpus

similar in the 2 sexes, with its length varied in the different phyletic lines.

Both of these features aré retained in Aedes (Nothoskusea) and relatively little

modified in Opifex.

The Culicinae do not offer unequivocal evidence for the number of waves of

dispersal in their population of New Zealand. However, the situation in the

Opifex-Aedes (Nothoskusea)-Aedes (Halaedes) complex suggests that at least 2

different dispersals, possibly 3, have occurred, the last one, involving aus-

tralis, being much more recent,

KEYS TO TRIBES AND GENERA

Adults

E, Spiraciiar Dristles PRCGOR) oi i. ec acee seca oar ke be ee ce

Spiracwiar Dristics absent... 6 4 gk a a ee

2(1). Wing with conspicuous bristles on lower surface of base of vein Sc (Culi-

SOUM) coos, 0 ce se 3 Culiseta

Wing without brigties on ‘lover surface of base ae vein Se ey

23,°M. aravropus).0) vie ke Pe ete oS. 2 e Miorisoelae

3(1). Postspiracular bristles: present (Aedini) .. 2.60 2 ow ON a. + 4

Postspiraculer bristles absent. 2. Sak a es a eee a

4(3). Scape of antenna strongly developed, visible as a distinct segment all

around at base of torus ... , °S Spa Opies

Scape of antenna a narrow ring, v visible aul at miesal articular point

UT OU ee re a de Ge oa eee

5(3). Claws with basal whitish-padlike pulvillus (Culicini). . . i) (es

Claws without basal whitish padlike pulvillus (Mansoniini) _ Coquillettidia

Male Genitalia

1. Paraproct of proctiger with conspicuous crown of numerous spicules

(Culiemi) .°: ei ie fe Culex

Paraproct of proctiger with oe 6 dpinelike teeth only. er PA Os a erry

2(1). Distal part of clasper irregular, with a more or less distinct subapical

lobe (Mansoniini) . ; . . «Coquillettidia

Distal part of clasper regula in shape, without any indication of a sub-

apies? lobé (Fee 2B es ee ee ee

3(2). Opisthophallic lobe strongly developed between the ane and procti-

ger (Sabethini; 23. M. argyropus) ...... . . . Maorigoeldia

Opisthophallie Jobe not differentiated.) go... ur eS

4(3). Apex of paraproct with 4 distinct teeth; mesal surface of sidepiece with-

out membrane (Culisetini; 14. C. téninbicd) ce se « « Culiseta

Apex of paraproct with only I tooth, simple or with small denticles; mesal

5(4).

3(2).

4(3).

5(4).

2(1).

3(2).

4(3).

(3).

Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

surface of sidepiece with more or less distinct membrane (Aedini) .

Cercal setae of 2 types, short distally, long proximally; sidepiece with-

out long bristles laterally and ventrally. . . ee, Opitex

Cercal setae all subequal and short; sidepiece with long bristles later-

Sry ancvenirally. Sie aie ae a ee et Aedes

Pupae

Trumpet without distinct erie apex specialized for piercing plant tis-

sues (Mansoniini) . er i ger: oa Som Leriaidia

Trumpet with distinct pinna: Pe re et ee ea ee

. Paddle with strongly produced apex and without apical hairs (Sabethini;

23. M. argyronus) <->. ‘ . . Maorigoeldia

Paddle rounded or emarginate apically, with at least hair i P developed

Hair 9-VIII on ventral surface, removed cephalad from caudolateral

margin (Culicini) . ; POE Oe Mea Ge Cue

Hair 9-VIII at posterolateral angle of sesment ae ee ee a. &

Paddle hair 2-P present in addition to 1-P but very small; hair 1-IX

present, simple (Culisetini; 14. C. tonnoiri) . : : See

Paddle hair 2-P not developed; hair 1-IX absent or » irregular (Aedini) .

Hair 9-VII simple, short, similar to hairs 9-II-V in shape and position

i a Page . Opifex

Hair 9-VII dorsal in position and markedly different from hairs 9-II-V

PODS Car as a es a ee ees . Aedes

Larvae

Median dorsal valve of siphon very long, fixed and with serrated inner

dorsal margin adapted for ions plant tissues eae

; . Coquillettidia

Median dorsal Valve at Biphen short, movable and never with serrated

aed Bedal Gia i eae a eae

Ventral brush of anal segment (4-X) with aie 1 = of hairs (Sabethini;

23. M. argyropus) . . . Maorigoeldia

Ventral brush of anal segment (4- -X) with Bi or more pairs of hairs. iS

Siphon with only 1 pair of subventral tufts (1-S) (Aedini). ...... 4

Siphon with several pairs of subventral tufts (1, 1a-S) . . 5

Metathoracic pleural group (9-12-T) very small, longest hairs less than

0.5 of long hairs in mesothoracic pleural group ..... . .Opifex

Metathoracic pleural group (9-12-T) at most moderately reduced, long-

est hair more than 0.5 of long hairs in mesothoracic pleural group

. Aedes

Siphon with a very small hair tuft (1-S) at the very base (Culisetini; 14.

Belkin: Culicidae of New Zealand 53

Cy tommetritewas tale a ae aw ee ee a a a ae ie Ca eeta

Siphon without such a differentiated hair at extreme base (Culicini) . .

. Culex

TRIBE CULICINI

Genus CULEX

1758. Culex Linnaeus, 1758:602. TYPE SPECIES: Culex pipiens Linnaeus, 1758,

Europe, selection of Latreille (1810:442).

For complete synonymy see Stone, Knight and Starcke (1959).

The species of Culex known to occur in New Zealand are readily recognized

from other Culicinae by the following combinations of characters. In the fe-

males, the tarsi are dark, the hindtibia has a conspicuous apical light spot, the

labium is light-scaled on the lower surface to the apical fourth, and the abdo-

men has basal light transverse bands on the tergites. In the males, the tarsi,

hindtibia and abdomen are the same as in the females, and the lower surface of

the palpus has a line of white scales on segment 4 and a basal spot of similar

scales on segment 5. In the pupae, hair9 on segment VIII is located on the ven-

tral surface a considerable distance cephalad of the posterolateral angle. The

siphon of the larvae is distinctive in the presence of Several subventral tufts

none of which is placed at the extreme base and the ventral brush is strongly

developed. A general account of the genus and tribe is given by Belkin (1962:

177-180).

SYSTEMATICS AND BIOGEOGRAPHY. The genus Culex is represented

in New Zealand by the introduced quinquefasciatus of the notorious pipiens com-

plex and 3 species of the endemic pervigilans complex. The two complexes

belong to the pipiens group (Belkin 1962, not Edwards 1932, 1941) of the subge-

nus Culex and are very similar in all stages but particularly in general exter-

nal adult characters, pupae and even male genitalia. There is a suggestion of

introgressive hybridization of quinquefasciatus and pervigilans in New Zealand.

C. quinquefasciatus has probably been introduced several times to New

Zealand on ships from the earliest days of sailing vessels to recent times. It

appears to be established only in the northern part of North Island (fig. 33) and

even here it is not very successful in competing with pervigilans and other na-

tive species, judging by our survey. While we found it breeding only in a few

large artificial containers and never in ground waters, Graham (1939:211) re-

ported it to breed "through the year in any place exposed to full sunlight and

holding water charged with decaying organic matter.'' Environmental conditions

do not seem to be particularly favorable for quinquefasciatus in New Zealand,

and this may be one of the reasons for its apparent hybridization with the native

pervigilans. However, during favorable periods quinquefasciatus may become

more abundant as reported by Graham.

Mattingly and Rageau (1958:241) placed pervigilans in their trifilatus sub-

group which included also iyengari Mattingly and Rageau, 1958 from New Cale-

donia, pacificus Edwards, 1916 from New Hebrides and Banks Islands, miracu-

losus Bonne-Wepster, 1937 from New Guinea, the Palaearctic torrentium Mar-

tini, 1925 and vagans Wiedemann, 1828, and the Ethiopian trifilatus Edwards,

1914 and tamsi Edwards, 1934 (Belkin 1962:fig. 84). In comparing the pattern

54 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

of distribution of the trifilatus subgroup to those of some species groups of

Neoculex, Ochlerotatus and Culiseta, Mattingly and Rageau made no comment

on the absence of this subgroup from Australia, where all the other groups are

represented. I suggest that the reason for this absence is that the trifilatus

subgroup is not really distinct from the pipiens complex and that the lineage of

the pervigilans complex, iyengari and pacificus is represented by 3 native spe-

cies in Australia, globocoxitus Dobrotworsky, 1953 and two forms currently

regarded as subspecies of pipiens, australicus Dobrotworsky and Drummond,

1953 and the "molestus" of these authors which is not molestus, Forskal, 1775.

I believe that the situation in Australia is probably analogous with that in New

Zealand in that introgressive hybridization may have occurred between the in-

troduced quinquefasciatus and the native australicus but that this has proceeded

farther with the formation of ''molestus'’ of Dobrotworsky and Drummond

(1953:132).

Edwards' tentative synonymy of torrentium with pervigilans (1932:210) lead

to the view that pervigilans may be nothing more than the European torrentium

introduced during the extensive whaling operations in the nineteenth century

(see Mattingly and Rageau). Now that 3 distinct species are known in the per-

vigilans complex in New Zealand this view should be discarded for it is most

unlikely that asteliae and rotoruae were formed in the recent past from an in-

troduced species.

Speciation in the pervigilans complex has taken place without distinct dif-

ferentiation in the male genitalia but with strong differentiation in the larva.

Two of the species are ecologically isolated, rotoruaein thermal waters, asteliae

in the leaf axils of terrestrialand epiphytic astelias. The dominant pervigilans

is a general ground water breeder which invades large artificial containers,

where it comes in contact with quinquefasciatus. C. pervigilans has a very

wide distribution with isolated populations on the Kermadec, Chatham and

Auckland islands. None of the isolated populations, with the exception of a

small one in Ocean Island in the Auckland group, is strikingly marked, and all

show a great deal of individual variation as do the populations on the main is-

lands of New Zealand. It seems probable that there has been considerable dis-

persal and intermixing of pervigilans populations through human transport and

that this species is clearly not prone to the formation of reproductively isolated

geographical populations.

Keys to Species

Females

1. Abdominal sternites II-VII with light scales only . .13. quinquefasciatus

Abdominal sternites II-VII with extensive dark scaling in the middle and

OO Te ee ee a we

2(1). Mesonotal scaling predominantly dark bronzy, fossa with numerous

short curved bronzy scales and without light scales. . . 11. rotoruae

Mesonotal scaling predominantly light coppery, golden or whitish, fossa

with light scales and at most with a few dark scales ....... 8

3(2). A conspicuous patch of curved bronzy scales between supraalar and pos-

terior dorsocentral bristles in front of level of wing . . .12. asteliae

2(1).

3(2).

a(1).

2(1).

Belkin: Culicidae of New Zealand 55

Scales on this patch usually light golden, not conspicuously differentiated

from other scales (not reliable) ©-. . 4 6.40. 4.0202 pervigilans

Males

Dark scales of sternites II-VII confined to median area . :

; ~ ee quinguefasciatus

Dark scales Of sternites II- vu more tee present on posterolater-

a) areas as -wWwellaS in (he middie. 1. ke ae et a a

Palpal segment 3 with no more than 10 long bristles in a single outer

row inthe distalhalf . . . . «++. ee, aeteline

Palpal segment 3 with at least 25 long bristles i. ina a double outer row in

Tie. CUSTAL (WOO Pde ke ee a a

Mesonotal scales predominantly bronzy on disc, with contrasting whit-

ish scales in lateral, supraalar and prescutellar areas . 11. rotoruae

Mesonotal scales predominantly light golden on disc, not markedly dark-

er Man CIBOWhENe. 6 fanaa heey, eo ee oe CO Ber visions

Male Genitalia

Basal sternal process of proctiger very short, weak and poorly pig-

mented; tergal arm of outer division of phallosome short and straight,

barely projecting beyond distal margin of inner division. :

wie crs. sainquefasciatus

Basal sternal process of proctiger long. strong, curved and heavily pig-

mented; tergal arm of outer division of phallosome very long, angled

distally and projecting well beyond distal margin of inner division .

: ; ; . 10. pervigilans; 11. rotoruae; 12. asteliae

Pupae

Pinna of trumpet strongly oblique, its maximum dimension at least 0. 4

of total trumpet length. .... ee . . 13. quinquefasciatus

Pinna more transverse, its maximum dimension less than 0.33 of total

brirhet length i Ee a oe at Gees -aiege Jue

Trumpet slender, index at least 8.0; hair 5-IV about 1.5 of length of

tergite V. .. : . . .12. asteliae

Trumpet moderately broad, andes ‘less than 6. 0: hair 5 IV shorter, usu-

en less than 1.25 of f length c of aa V. : ;

eee ae .10. pervigilans: 11. rotoruae

Larvae

Ventral brush with 7 pairs of hairs; siphon usually with 5 pairs of tufts

all below level of pecten and more or lessinline. . . . I1. rotoruae

Ventral brush with 6 pairs of hairs; siphon usually with 4 pairs of tufts,

l or 2 pairs out of line above level of pecten: ... 5 2

Two pairs of siphonal tufts clearly above level of pecten; prothoracic

hairs 7, 8-P both single; saddle hair 1-X usually double. .12. asteliae

Usually only one pair of siphonal tufts above level of pecten (out of line)

06 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

or all hairs more or less in line; prothoracic hairs 7,8-P usually

both double; saddle hair 1-X usually single .. . 2... 5... 8

3(2). Hair 1 on abdominal segments III-V usually at least 4-branched; mental

plate usually with less than 9 teeth on each side of apical tooth; sub-

apical siphonal tuft usually only slightly out of line; siphon relatively

long, index usually more than 5.0 .. ios jee + LO. pervigilans

Hair 1 on abdominal segments III-V usually single or double; mental

plate usually with at least 10 teeth on each side of apical tooth; sub-

apical siphonal tuft usually conspicuously out of line; siphon relative-

ly short, index usually less than 5.0. . .. . . .13. quinquefasciatus

10. Culex (C.) pervigilans Bergroth

Figs. 14, 30, 31

1889. Culex pervigilans Bergroth 1889:295. TYPE: Holotype 2, Greymouth,

Westland, New Zealand [Location unknown].

Culex (C.) pervigilans of Edwards (1924:396; 1932:209); Graham (1929:216-227;

1939:211); Taylor (1934:24); Lee (1944:109); Miller (1950:44); Miller and

Phillipps (1952:19); Mattingly and Rageau (1958:241); Stone, Knight and

Starke (1959); Dumbleton (1962:24, 25; 1965:142-143); Belkin (1962:190-191);

Nye and McGregor (1965:4-7).

Culex (C.) sp. 3, Auckland Island form of Belkin (1962:191).

Culex sp. of Harrison (1956:211).

FEMALE (Graham 1929:figs. 25-32; Miller and Phillipps 1952:figs. 32-34).

Wing: 4.5 mm. Proboscis: 2.1mm. Abdomen: about 3.0 mm. In general sim-

ilar to members of the pipiens complex from which it is distinguished in New

Zealand by presence of dark scales in posterolateral areas and middle of ab-

dominal sternites and other characters mentioned under quinquefasciatus. Ex-

tremely variable in most features and very similar to other members of per-

vigilans complex from which it cannot be always separated. General coloration

of integument very dark, usually blackish brown on mesonotum and head; mes-

onotal scales largely elongate and straight, usually pale and contrasting sharp-

ly with mesonotal integument. Head: Decumbent scales whitish, creamy to

golden; erect scales similar in color to decumbent in the middle and in front,

a variable patch of darker scales posterolaterally, these sometimes blackish.

Labium conspicuously pale-scaled ventrally usually to apical 0.25, sometimes

pale-scaled dorsally near middle. Palpus usually with whitish, creamy or

golden scales on segments 3 and 4; these scales sometimes very conspicuous.

Flagellar segment 1 usually with distinct patch of pale scales as on palpus.

Thorax: Mesonotal scales usually predominantly whitish, creamy or golden,

sometimes a more or less conspicuous supraalar patch of shorter, more

curved, light bronzy scales. Apn and ppn scales as on mesonotum; psp usually

without hairs or scales; stp and mep scales usually pure white, rarely creamy

or golden (Auckland Is. populations). Legs: Femora usually predominantly

dark-scaled on anterior surface, rarely hindfemur indistinctly paler on ante-

rior surface ventrally, ventral and posterior light scaling restricted; knee

spots and apical hindtibial spot usually large and conspicuous, usually whitish;

lower surface of tibiae and basal tarsal segments not distinctly pale; midcoxa,

hindcoxa and extreme base of all femora with light scales; remainder of legs

Belkin: Culicidae of New Zealand 57

dark-scaled. Claws all without teeth and all with short fine spicules on outer

surface of base. Haltere: Upper part of stem and all of knob with pale brown

scales. Abdomen: Tergite I with small median apical patch of dark scales,

rarely with a few light scales; tergites II-VII with basal transverse light bands

which are lengthened caudad at lateral ends progressively more from III to VII,

median part straight or widened in the middle particularly on III,1V,V, light

scaling usually whitish; sternites with varied amount of dark scaling but al-

ways with at least posterolateral angles and basal median areas dark-scaled

on distal segments, pale scaling usually whitish.

MALE (Graham 1929:fig.34). Essentially as in the female. Palpus ex-

ceeding proboscis by at least full length of segment 5, sometimes more; seg-

ment 4 with white-scaled line ventrally from base to apex, segment 5 with small

basal ventral patch of white scales, segment 3 with lighter scales dorsolateral-

ly; segment 3 usually with at least 25 very long external ventral hairs in a

double row in the distal half to two-thirds; segments 4 and 5 with dense vesti-

ture of very long ventrolateral hairs on both sides. Labium without ventral pale

streak, entirely dark except for very inconspicuous lighter area around false

joint.

MALE GENITALIA (fig. 14; Graham 1929:fig.33; Nye and McGregor 1964:

figs. 3a,b). As figured here and as described by Belkin (1962:190); apparently

indistinguishable from other members of the complex. Subapical lobe of side-

piece with 1-3 setae in group d-f not 1 as stated by Belkin (loc. cit.). Inner

division of phallosome short, bent as in pipiens complex; lateral arm of outer

division strongly developed, in dorsal aspect appearing angled and bearing fine

spicules internally; tergal arms very long, in dorsal aspect divergent, angled

mesad distally and extending at least to level of apex of proctiger. Basal ster-

nal process of proctiger very long, sharply curved ventrad and strongly pig-

mented.

PUPA (Belkin 1962:fig.90; Graham 1929:figs. 48,49). Abdomen: 3.7 mm.

Trumpet: 0.75 mm. Paddle: 1.05 mm. As figured and described by Belkin

(1962:190); very similar to quinquefasciatus and asteliae, and indistinguishable

from rotoruae. Trumpet relatively short and broad, index usually less than

6.0; pinna usually less than 0.25 of trumpet length; usually darkened at base

and apex and lighter in the middle. Hair 5-IV usually shorter than tergite V.

Both paddle hairs usually present.

LARVA (Belkin 1962:fig.91; Graham 1929:figs. 37-47; Lee 1944:plate 82;

Miller and Phillipps 1952:fig. 36; Nye and McGregor 1964:fig.4a). Head: 0.92

mm. Siphon: 1.55mm. Saddle: 0.46 mm. As figured and described by Bel-

kin (1962:190); in general similar to asteliae, rotoruae and quinquefasciatus; ex-

tremely variable but readily distinguished from the above by the diagnostic

characters in the key and the following features. Head capsule uniformly mod-

erately to strongly pigmented; mental plate with 6-8 coarse teeth on each side

of median apical tooth; aulaeum very conspicuous and projecting. Antenna long

and very slender, usually at least 0.7 of head length; usually uniformly darkly

pigmented; spicules strong, usually extending distad of hair 1. Prothoracic

hairs 4,7 and 8 usually all double, rarely triple. Abdominal hair 1-III-V

usually 3, 4b; hair 6-III-VI usually double or triple. Siphon usually uniformly

moderately or darkly pigmented, sometimes darkened at base and apex; ex-

tremely variable in length but index usually about 5.5-7.0; usually 4 subventral

tufts (1, la-S), subapical slightly to distinctly out of line, proximal usually dis-

tad of pecten, all longer than width of siphon but very variable in length and

number of branches. Saddle usually moderately to strongly pigmented, spicu-

58 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

lation usually inconspicuous; hair 1 usually single. Ventral brush usually with

6 pairs of hairs. Anal gills subequal; variable in length, usually distinctly long-

er than saddle.

SYSTEMATICS. As interpreted here, C. pervigilans is readily separated

from the other 2 members of its complex, rotoruae and asteliae, in the lar-

val stage by features of the siphon and anal segment, from asteliae in the pupal

stage by the shape of the trumpet and from rotoruae in the adults by the meso-

notal scaling. Its pupa is apparently indistinguishable from rotoruae and the

adults cannot be separated with certainty from asteliae. The male genitalia ap-

pear to be very Similar in all 3 species, but with more detailed study it may

be possible to separate asteliae from the other 2 species of the pervigilans

complex.

C. pervigilans is the dominant, commonest and most widespread culicid of

the New Zealand region. It exhibits a great deal of variation in nearly all fea-

tures of all stages even in the diagnostic ones, although in the latter there is no

overlap with either rotoruae or asteliae. There appears to be nearly as much

intrapopulation as interpopulation variation in most characters, suggesting that

considerable hybridization between originally isolated populations is occur-

ring at the present and probably has been going on for some time. As indicated

under quinquefasciatus thereis a suggestionof introgressionof some pervigilans

features in the populations near Auckland, and it is possible that some intro-

gression of quinquefasciatus features has occurred in pervigilans populations

in other areas.

There is considerable geographical differentiation in pervigilans in the

isolated insular populations. It is very likely that considerable mixing in these

populations has occurred as a result of transport of breeding populations from

place to place on sailing vessels and motor ships since early in the last centu-

ry. Itis, of course, possible also that some of these insular populations were

originally established through such human agency. In none of the remote insu-

lar populations is there differentiation in the larval stage approaching that found

in the ecologically separated species of the complex on the main islands of New

Zealand, even at the two extremes, Kermadec and Auckland islands, where en-

vironmental conditions must be very different.

The Kermadec population appears to be quite different from the other re-

mote insular populations but unfortunately the material is very limited. There

are only 5 fourth-instar larvae in the single larval collection. These have a

general facies quite distinct from the New Zealand populations. The pigmen-

tation of the head capsule, antenna, all hairs, siphon and anal saddle is much

lighter; the hairs are generally weaker; the anal gills are distinctly shorter

than the anal saddle; hair 6-III-V is usually triple, and prothoracic hairs 3, 7,

8-P are sometimes triple; the teeth of the mental plate are 6 or 7; the siphon is

long and slender and without marked tapering distad, its index is about 6.0-7.0;

the position of the siphonal tufts (1, la-S) tends to be irregular, the basal one

is sometimes with the pecten and the preapical one tends to be strongly dis-

placed dorsad. There is considerable variation in nearly all these features and

all the variations encountered are frequently found in the populations from the

main islands of New Zealand but not in exactly the Same combination in a

single population. The mesonotal scaling of the adults tends to be niore yellow-

ish and the mesonotal bristles shorter than in the typical main New Zealand

populations. The male genitalia show no unusual characters; there is only 1 |

seta in group d-f and 8-10 bristles on the lobe of tergite IX. I see no reason

for considering this as anything but a small isolated population of pervigilans

Belkin: Culicidae of New Zealand 59

adapted to subtropical insular conditions.

The Chatham Islands population is the most isolated of all pervigilans pop-

ulations, this island group being about 500 miles from Cape Pallister at the

southeast end of NorthIsland. In the adequate sample of larvae there is more

variation than is usually found in individual New Zealand collections, as would

be expected in smaller isolated insular populations, but all of the features of

the Chatham population fall within the range of variation of pervigilans on the

main islands. The general facies of these larvae is quite different from that of

the Kermadec population and appears more similar to that of the Auckland Is-

lands populations but is not markedly different from the main island populations

although it is more or less distinct. The pigmentation of the sclerotized parts

and hairs is moderate to strong; the larger hairs have strong branches; the

anal gills are usually only slightly longer than the anal saddle; the antenna

tends to be shorter than in New Zealand, the siphon is shorter (index 5.0-5. 5);

the proximal siphonal tuft (1-S) tends to be within the pecten and the 2 distal

tufts usually have more branches; prothoracic hairs 4, 7, 8-P are frequently

triple as is abdominal hair 6-III-V. Except for 2 males the few reared adults

are in alcohol and it is not possible to determine accurately the color of the

mesonotal scales. The male genitalia are essentially similar to those of the

main island populations; there are 1 or 2 setae in group d-f and 6-12 bristles

on the lobe of tergite IX. Inthe male, segment 3 of the palpus has an indistinct

line of pale scales dorsolaterally above the long external bristles and the labi-

um an indistinct pale ring around the false joint. In my opinion, the Chatham

Islands population is not sufficiently differentiated to be recognized as a sepa-

rate taxonomic entity. :

The Auckland Islands populations are of considerable interest because they

are found near the southern known limit of occurrence of the family. The lar-

vae in the 3 collections are quite similar and are not markedly different

from the Chatham Island population except that they usually have more branches

in abdominal hair 6-III-V (5-7), prothoracic hairs 4, 7,8-P are more frequent-

ly triple and may have even 4 or 5 branches, and the siphon is a little longer.

As in the other isolated insular populations there is a great deal of individual

variation in larval characters. The sample of adults is very small but very in-

teresting because of apparent striking differences between the Ranui population

on the main Auckland Island and the population on the adjacent very small

Ocean Island. The Ranui adults are of normal size and coloration and the

lobe of tergite IX bears 6-9 bristles. The Ocean Island adults are the largest

I have seen (wing length of female, 6 mm), the light scaling of the head and

thorax is distinctly yellowish and the lobe of tergite IX of the male genitalia

bears 16-18 bristles. Males of both populations have a distinct line of whitish

scales in the middle of segment 3 of the palpus above the long external bristles;

Ranui males have an indistinct narrow pale ring around the false joint of the

labium and in the Ocean Island males this light ring is much larger and more

conspicuous; one male from Ranui has 2 setae in group d-f, all the others have

only one. In females of both populations the light ventral scaling of the labium

extends dorsad to form an indefinite broad pale ring which is larger and more

conspicuous in the Ocean Island population. If any population of pervigilans

deserves recognition as a separate taxonomic entity, it is the Ocean Island one.

However, before this is done, more material is needed to evaluate the puzzling

situation of the presence of 2 apparently distinct populations in the Auckland Is-

lands. Several possibilities exist to account for these different populations in-

cluding the presence of an indigenous form and the introduction of one or more

60 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

populations at different times in the past.

I have made no attempt to analyze critically the populations of pervigilans

from the main islands of New Zealand and the adjacent small islands. This

would require a Special study with more material than I have available. How-

ever, a Superficial examination reveals a great deal of variation in all stages

without any clear indication of geographical differentiation. In the larva partic-

ularly, there is bewildering variation in nearly all features: in the length and

pigmentation of the antenna; in the pigmentation of the head capsule and the

branching of head hairs; in the branching and character of the larger thoracic

and abdominal hairs; in the length and shape of the siphon, the number of pec-

ten teeth, and the branching, distribution and character of the siphonal tufts;

and in the length of the anal gills. However, there is no significant overlap in

any of the characters which are diagnostic of rotoruae and asteliae in the larval

stage; the antenna is never as short as in rotoruae and the ventral brush never

has 7 pairs of hairs as in this species; the siphonis never as long as in asteliae

and never has the characteristic arrangement and branching of the siphonal

tufts of this species. In the pupa, the pinna of the trumpet is extremely variable

and sometimes approaches the condition in quinquefasciatus. In the adults, the

mesonotal and head scaling also show a great deal of variation, some of which

is probably due to the partially teneral condition of the reared specimens. In

the male genitalia the most conspicuous variation is found in the number of

setae in group d-f (1-3) and the number of bristles on the lobe of tergite IX (6-

13); in the case of the latter, however, the number seems to be relatively con-

stant within a given population.

BIONOMICS. C. pervigilans is the dominant and most widespread culicid

of the New Zealand area, probably because of its wide tolerance of environmen-

tal conditions as shown by its very wide range of breeding sites. It has been

reported by Graham (1929:221, 226) to breed in all types of ground waters,

still and with moderate flow, clean and contaminated, fresh and with as much

as 75% sea water, and from a multitude of small and large artificial containers.

In our field work we found pervigilans most commonly in natural breeding sites

(stream margins, springs, pools, ponds and swamps) and in ditches and drains,

and much less frequently in various types of artificial containers (concrete cis-

terns, metal tanks, drums and pans). This species has been found associated

with all other culicids breeding in artificial containers and ground waters ex-

cept O. fuscus and A. chathamicus. Breeding continues throughout the year at

least around Auckland (Graham 1929:226).

C. pervigilans is apparently a serious nocturnal domestic pest in some

years both in cities and rural areas (Graham 1929:219) but does not seem to

attack man readily out of doors or in the field. All of our adult collections

were made in buildings. The natural hosts of pervigilans are probably birds.

Ross, Austin et al (1963) have isolated viral agents from pervigilans.

DISTRIBUTION (figs. 30,31). Most widely distributed culicid of the New

Zealand area from Kermadec Islands to Auckland Islands and Chatham Islands.

Material examined: 4180 specimens; 327 0, 364 2, 2805 larvae, 684 pupae; 170

individual rearings (82 larval, 66 pupal, 22 incomplete).

KERMADEC ISLANDS. Raoul Island: Station cistern, 28 Sept 1962, G.A.

Samuelson (89), 63 L[BISH]. Bell's Ravine, elev. 75m, Malaise trap, G.A.

Samuelson, 7-12 Sept 1962, 19, 13-14 Sept 1962, 179 [BISH]. N Slope Ridge,

elev. 150 m, light trap, 6-11 Oct 1962, G.A. Samuelson, 1 o [BISH]. Locali-

ty not specified, 9 Oct 1908, 3 o° [CANT]; Aug 1908, 1¢%, [WELL]. |

NEW ZEALAND. North Auckland: Waipapakauri, 26 June 1944, G. L.

Belkin: Culicidae of New Zealand 61

Bissett (371), 1 L[NELS]. Awanui, 16 Feb 1944, W.J. Phillipps (43), 1 L

[NELS]. Whangaroa, 12[WELL]. Kaeo, 3 miS on route 10, 20 Dec 1963,

1963, BS (NZ 39), 1 lpo& (39-101), 10 L[UCLA]. Mangamuka Gorge, 2 mi N

of Soda Spring, 21 Dec 1963, BS (NZ 41), 71 L[UCLA]. Ngawha, 16 Feb 1944,

W.J. Phillipps (46), 3 L[NELS]. Opua, 3 mi S of turnoff on Kawakawa- Paihia

road, 20 Dec 1963, BS (NZ 38), 2 L[UCLA]. Waipoua forest, 19 Mar 1945,

W.J. Phillipps (642), 3 L[NELS]. Whangarei, 8 Jan 1919, D. Miller, 3

[BMNH]; 10 Jan 1919, D. Miller, 1%, 22[BMNH]. Ruakaka, 1 mi S on high-

way 1, 20 Dec 1963, BS (NZ 36), 4 1po (36-101, 106, 108, 109), 1 1p? (36-110),

1 po’ (36-102), 3 p? (36-103-105), 2 1p (36-107, 112), 10%, 29, 111 L, 12P

[UCLA]. HenIs (Taranga), 1925, 1“ [WELL]. Wellsford, elev. 100m, 31

May 1964, F (NZ 188), 4L[UCLA]. Martins Bay, Mahurangi Heads, 26 Dec

1964, F (NZ 196), 1 L[UCLA]. Parakai, 18 July 1918, D. Miller, 10, 29

[BMNH]. Hobsonville, 25 Oct 1944, G.L. Bissett (419), 1 L[NELS]. Bethells

Road Swamp, 9 May 1964, F (NZ 184), 1 L[UCLA]; 10 Jan 1965, F (NZ 200B),

1L, 2 P[UCLA]; 12 Jan 1965, F (NZ 202), 1 L[UCLA]. Waitakere Ranges,

Quarry tributary of Waitakere River, 18 Dec 1963, BS (NZ 35), 4 L [UCLA];

0.25 mi S from main turnoff to Karekare from Piha Road, 14 Jan 1964, S (NZ

97), 70 L, 1 P[UCLA]; 2 Nov 1944, G.L. Bissett (442), 10°, 19 [NELS].

Oratia, DSIR Field Station, 22 Dec 1963, BS (NZ 45), 7 L [UCLA]; 30 Dec 1963,

S (NZ 66), 7 L[UCLA]. Titirangi Beach, 30 Dec 1963, BF(NZ 64), 1L[UCLA].

Titirangi, 9 Jan 1946, G.L. Bissett (831), 3 L[NELS]; 18 Mar 1946, G. L.

Bissett (891), 19 [NELS]; swamp in field near forest, 1 July 1946, G.L. Bis-

sett (975), 1 L[NELS]. Takapuna, 1943, W.J. Phillipps, 1 o [WELL]. Devon-

port, Shoal Bay, 18 Jan 1917, D. Miller, 1 o [BMNH]. Northcote, Clarence

Road, 30 Sept 1963, S(NZ 9), 1¢%, 33 L, 4 P[UCLA]. Auckland, Arch Hill,

catch basin, 3rd Ave, 26 Mar 1919, D. Miller, 1 o, 1 ?2[BMNH]; Avondale, 23

Nov 1965, F (NZ 216B), 1? [UCLA]; Cabbage Tree Swamp, 26 Mar 1919, D.

Miller, 1 9[BMNH]; Ellerslie, gully swamp, race course, 24 Mar 1919, D.

Miller, 10, 19[BMNH]; Kaitaria Swamp, 2 Feb 1917, D. Miller, 1¢ [BMNH];

Kohimarama, 29 Apr 1946, G. L. Bissett (926), 2 L[NELS]; Mt. Albert, 1916-

1925, A.E. Brooks, 7%, 3?[BMNH]; Mt. Wellington, 14 June 1945, G.L.

Bissett (736), 2°, 19 [NELS]; Newton, 22 Feb 1944, Francis, 30, 119

[WELL]; Owairaka, 2 July 1946, G.L. Bissett (976), 2%, 2 9 [NELS]; Penrose,

18 May 1944, G.L. Bissett (328), 1 L[NELS], 15 July 1946, G.L. Bissett (990),

4 L[NELS]; Three Kings, 11 July 1946, G. L. Bissett (982), 6 L [NELS]; West-

mere, 17 Nov 1944, G.L. Bissett (464), 1 L[NELS], 11 Jan 1946, G.L. Bis-

sett (836), 2 L[NELS]. Auckland, locality unspecified, 27 July 1923, O.H.

Swezey, 19[BISH]; 18 Dec 1941, T.R. Rabone, 1 0, 1 9 [AUCK]; ex spring,

20 Jan 1950, K. Wise, 19 [AUCK]. Mangere, 7 Jan 1965, F (NZ 201), 19, 1P

[UCLA]; 26 Oct 1950, K. Wise, 19 [AUCK]; Oct 1950, K. Wise, 1 9 [AUCK].

Howick, 24 May 1917, A.E. Brookes, 10°, 29[BMNH]. Hunua Range, 1 Jan

1916, A.E. Brookes, 1¢[BMNH]. Pukehoke, sewage oxidation plant, 25 May

1964, F (NZ 186), 1 lp? (186-112), 1 p? (186-111) [UCLA]. Tuakau, 9 June

1964, F (NZ 190), 1 1po (190-121) [UCLA].

Little Barrier Island: Near Ranger's house, 9 Oct 1963, S(NZ 10), 16 L

[UCLA]. Near Ranger's chicken house, 6 Mar 1964, S (NZ 129), 1 lpo (129-

104), 1 1p? (129-103), 3 po (129-102, 109, 112), 7 p? (129-108, 110, 111, 113-

117), 50, 149, 38 L, 41 P[UCLA]. Small stream between Te Waikohare and

Tirikikawa streams, 9 Oct 1963, S(NZ 11), 3 1p (11-102, 106,107), 41p9

(11-101, 103-105), 2 1p (11-108, 110), 30°, 38 L, 3 P[UCLA]; 6 Mar 1964, S

(NZ 128), 1 po (128-118), 3 p? (138-101, 102,115), 41%, 149, 112 L, 80 P

62 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

[UCLA]. Summit Track, elev. 267 m, 9 Oct 1963, S(NZ 12), 1 L[UCLA]. Be-

tween Lamb Bay creek and Awaroa stream, 31 Oct 1963, S (NZ 20), 4c, 59,

19 L, 8 P[UCLA]. Awaroa stream, 0.5 mi upstream, 31 Oct 1963, S (NZ 22A,

B), 3%, 10 L, 5 P[UCLA]; 0.25 mi upstream, 31 Oct 1963, S (NZ 23A, B),

6%, 39, 24L, 10 P[UCLA]. Waipawa stream, 6 Mar 1964, S (NZ 130), 3 1p?

(130-112-114), 1 po’ (130-109), 6 p? (130-110, 111, 115-118), 170°, 159, 73 L,

45 P[UCLA]. Tirikikawa stream, 8 Mar 1964, S (NZ 131), 1 L [UCLA].

Great Barrier Island: Shoal Bay, 25 Oct 1963, S (NZ 16), 1 1p (16-105),

14 L[UCLA]. Te Wairere stream, 26 Oct 1963, S(NZ 17), 3 L, 1 P [UCLA].

Locality not specified, 16 Feb 1919, D. Miller, 1 o% [BMNH].

Cuvier Island: 13 Mar 1963, S (NZ 136), 1 L [UCLA].

South Auckland: Coromandel, 25 Apr 1964, N (NZ 220), 19 L[UCLA]. Te

Puke, Prew Orchard, 31 Mar 1964, S (NZ 142), 5 L[UCLA]. Karapiro, 25 Jan

1946, A.W. Jamison (852), 30°, 39 [NELS]. Te Kuiti, 18 Dec 1944, G. Milne

(500), 10’, 19; 30 Jan 1946, J. Beattie (853), 20, 19 [NELS].

Gisborne: Waioeka Gorge, 19 Nov 1963, S (NZ 29), 12, 18 L, 2 P[UCLA].

Ormond, 17 Nov 1963, S (NZ 28), 40°, 32, 18 L, 7 P[UCLA]; Waihirere Do-

main, 28 Mar 1964, S (NZ 140), 3 L[UCLA]. Hexton, 26 Dec 1963, BS (NZ

50), 3 1p (50-101-103), 2 po (50-106, 110), 5 p? (50-104, 105, 107-109), 1 9,

25 L, 9 P[UCLA]. Mokau Falls, Urewera National Park, 27 Dec 1963, BS (NZ

55), 21po' (55-101, 102), 1 1p (50-103), 20°, 19, 54.L, 9 P[UCLA]. Lake

Waikaremoana, 27 Jan 1957, E.N. Marks (9,10), 20, 12, 4 L[UCLA].

Hawke's Bay: Cape Kidnappers, 16 Nov 1963, S (NZ 27), 5 1po (27-101,

104, 106, 109, 110), 2 1p? (50-102, 107), 3 1p (50-103, 105,108), 19, 14L, 4P

[UCLA]. |

Wellington: Tokaanu, Dec 1943, W.J. Phillipps, 2? [WELL]. Pokaka, 10

Jan 1964, BFS (NZ 86), 1 lpo‘ (86-101), 1 1p (86-107), 1 po (86-104), 14 L

[UCLA]. Ohakune, Rangataua Reserve lake, 9 Jan 1964, BFS (NZ 84), 168 L

[UCLA]; pond at bend of Lake road, 9,10 Jan 1964, BFS (NZ 85), 1 L[UCLA],

26 Jan 1964, F (NZ 174), 4L, 5P [UCLA], 28 Jan 1964, F (NZ 176), 39

[UCLA]; locality not specified, 1°, Jaly-Aue 1922, 7. R. Harris [BMNH], 1-9

Apr 1923, T.R. Harris, 1 oc [BMNH]. Kaitawa, Nov 1922, 1A [WELL]. Otaki,

30 Dec 1943, W.J. Phillipps (184), 4L, 1P [NELS]. Masterton, 10 Jan 1945,

F. Ellett (515), 20°, 29[NELS]. Kapiti Island, 1 Oct 1928, 19[WELL]. Pa-

raparaumu, 1944, W.J. Phillipps, 3%, 32 [WELL]. Lower Hutt, 15 Nov 1945,

W.J. Phillipps (827), 3c [NELS]. Featherston, 18 Jan 1945, F. Ellett (510),

19 [NELS]. Castle Pt., Wairarapa, 7 Dec 1945, F. Ellett and W.J. Phillipps

(829), 5L, 2 P [NELS]. Wellington, 24 Apr 1924, 1 9 [WELL], 6 Jan 1929,

J.T. Salmon, 2 0° [WELL], 29 Nov 1943, 1 o [WELL], 4 Jan 1944, W.J. Phil-

lipps, 1c, 29[WELL], 22 Feb 1944, W.J. Phillipps, 30°, 49 [WELL]; Kel-

burn, 29 Feb 1944, W.J. Phillipps, 5 ° [WELL]; Ohariu Valley, 7 Feb 1944,

W.J. Phillipps (16a), 1 L[NELS]; Rocky Bay, 14 Dec 1943, 1 o [WELL]; Wilton

Bush, Feb 1922, G.V. Hudson, 12 [BMNH]; Wilton Road, Jan 1944, 20°, 19

[WELL].

Nelson: Canaan Beech Forest, Harwood's Hole, 4 Jan 1964, BG (NZ 77),

to. 28. 1 P UCLA: Harwood's Hole track pond, 4 Jan 1964, BG (NZ 78),

1 lpo (78-101), 9 Ip? (78-102-110), 12 ¢, 72, 105 L, 21 Pp [UCLA]. North

Branch Riwaka River, 4 Jan 1964, BG (NZ 75), aL, [UCLA]. Wakapuaka, 12

Mar 1964, S (NZ 167), 1 L [UCLA]. Nelson, 3 Jan 1964, BG (NZ 73), 8¢,

42, 29 L, 11 P TUCLA], 16 Mar 1923, A.L. Tonnoir, 2 o, 1 9 [NELS], 1°L

(696) [NELS]; Cawthron Institute, 24 Jan 1945, W.J. Phillipps (548), 1 L [NELS];

Blick's Valley, 12 Mar 1964, S (NZ 164), 2 L, 1 P[UCLA]; Maitai River, junc-

Belkin: Culicidae of New Zealand . 63

tion of Sclanders stream, 11 Feb 1964, S (NZ 125), 3 L[UCLA], south Branch,

11 Feb 1964, S (NZ 126), 32 P[UCLA], near Pole Ford Bridge, 14 Mar 1964,

S(NZ 170), 8 L, 2 P[UCLA]; North Road, 16 Nov 1943, 1? [WELL]; Reser-

voir, 12 Mar 1964, S (NZ 163), 2 1p? (163-101, 102), 2.0, 126L, 44 P [UCLA];

Tahunanui, 10 Mar 1964, S (NZ 157), 30%, 50 L, 8 P [UCLA]. Stoke, 11 Mar

1964, S (NZ 160), 13 L, 12 P [UCLA]. Aniseed Valley, 21 Mar 1922, A. L.

Tonnoir, 1¢ [NELS]. Poorman's Valley, 30 Nov 1944, E.S. Gourlay (472),

1A [NELS]. Wairoa Valley, 23 Jan 1945, B.B. Given (546), 2 L [NELS].

Howard Junction, 13 Mar 1964, S (NZ 169), 22 L, 3 P[UCLA]. Kawatiri Junc-

tion, 9 Feb 1964, S (NZ 120), 1 1p? (120-106), 2 po’ (120-101, 102), 1 p? (120-

104), 21p (120-105, 107), 160°, 5 9, 68 L, 46 P [UCLA]. New Creek Road,

between Murchison and Kawatiri Junction, 8 Feb 1964, S (NZ 116), 46L [UCLA].

Gowanbridge, 9 Feb 1964, S (NZ 122), 12 L[UCLA]. Gowan River, 9 Feb 1964,

S (NZ 121), 1 po (121-103), 10 L [UCLA]. Lake Rotoroa, 21 May 1923, C.L.

Edwards, 1 [BMNH].

Marlborough: Blenheim, 18 Oct 1945, W.J. Phillipps (823), 19 L, 1 P

[NELS]. Kaikoura, Feb 1922, A.L. Tonnoir, 1 o [NELS].

Westland: Greymouth, date and collector not specified, holotype 9 | Berg-

roth 1889:295, location unknown]. Hercules Mountain, 7 Feb 1964, S (NZ 115),

11 L[UCLA]. Whataroa, 7 Feb 1964, S (NZ 114), 1 po (114-102), 3 p? (114-103,

106,107), 13 L[UCLA]. Waiho, 7 Feb 1964, S (NZ 113), 38 L[UCLA]. Local-

ities not specified, A[J.S. Pillai 1966:25].

Canterbury: Kaiapoi, 28 Aug 1963, S(NZ 3B), 39 [UCLA], 21 Jan 1964,

S (NZ 99), 1%, 39[UCLA]. New Brighton, Nov 1943, S. Lindsay, 1c, 39

[WELL]. Christchurch, F.W. Hutton (90), 1, 89 [BMNH], 21 July 1899,

R. L.. Scott, 30°, 49 [BMNH], 17, 18 Feb 1922, A.L. Tonnoir, 10°, 1 9 [NELS].

Akaroa, Dec 1945, D. Miller (830), 10 L[NELS]. Timaru, 24 Jan1964, S (NZ

100), 1 1po (100-107), 2 1p (100-102, 109), 2 pot (100-108, 112), 3 p? (100-

106, 111,113), 11p (100-105), 271 L, 22 P[UCLA]. Makikihi, 25 Jan 1964, S

(NZ 101), 29 L[UCLA]. |

Otago: Kinloch, 11 Jan 1957, E.N. Marks (21, 22), 10°, 19 [UCLA].

Frankton, L [Nye and McGregor 1964:7]. Middlemarch, L [Nye and McGregor

1964:7]. Dunedin, Leith Valley, 1 Aug 1922, G. Howes, 1 9 [BMNH]; Otago

Museum grounds, L [Nye and McGregor 1964:7 ]. Otago Peninsula, Hooper's

Inlet, 2 Jan 1964, BN (NZ 71), 1 1p (71-101), 11p (71-102), 20, 19, 76L,

3 P [UCLA]; Allan's Beach, Doctor's Point, Hooper's Inlet, L [Nye and Mc-

Gregor 1964:7]. Taeri Plain, L [Nye and McGregor 1964:7]. Catlins District,

19 Jan 1957, E.N. Marks (26), 1o, 19, 4L, 2 P[UCLA], L [Nye and Mc-

Gregor 1964:7].

Southland: Te Anau, 2 Feb 1964, S(NZ 110), 1 1p? (110-104), 1 po (110-

102), 1 p2 (110-101), 1 1p (110-103), 29 L, 4 P [UCLA]. Between Invercargill

and Bluff, 28 Jan 1964, S (NZ 107, 108), 11po (107-101), 2 1p? (107-103, 104),

1 po’ (107-105), 1 p9 (107-102), 1c, 19, 27 L, 9 P (107), 53 L, 1 P (108)

[UCLA]. Bluff, 26 Jan 1964, S(NZ 104), 2 1po (104-113, 183), 2 lp? (104-111,

112), 2 pot (104-182, 184), 1 p? (104-181) [UCLA].

Stewart Island: Oban, road to Observation Rock, 27 Jan 1964, S (NZ 105),

2 1po (105-104, 111), 4 1p (105-105-107, 110), 5 po (105-102, 108, 109, 112,

113), 1 pQ (105-101), 2 1p (105-103, 114), 50, 309, 66 L, 123 P[UCLA].

Golden Bay, 27 Jan 1964, S(NZ 106), 1 1p (106-108), 4 1p (106-102, 104,

106,107), 1 po (106-101), 1 p? (106-103), 10 L[UCLA]. |

New Zealand, locality unspecified or unknown: Locality not specified, 15-

22 July 1923, T.R. Harris, 1 o [BMNH]; 1 9 [WELL]; 1957, E.N. Marks (1, 2),

64 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

6 L, 1 P[UCLA]; 1921, D. Miller, 1 [UCLA]; 2 L (345), 2 (488), 22(506),

12 (507), 13 L (617), 30, 39 (901), 16, 29 (917) [NELS]. Waituhi(?) forest,

26 Mar 1945, W.J. Phillipps (636), 2 L[NELS]. Tutara, Urewera, 26 Mar

1945, W.J. Phillipps (661), 1 L [NELS].

CHATHAM ISLANDS. Flying boat base, 26 Feb 1964, S(NZ 151), 21p¢

(151-103, 104), 1 1p (151-106), 1 1p (151-105), 11 L[UCLA]. Owenga, 26 Feb

1964, S(NZ 152), 3 lpo’ (152-101, 102,107), 2 1p@ (152-105, 106), 2 1p (152-

103, 104), 98 L[UCLA]. Waitangi, 29 Feb 1964, S (NZ 154), 3 p (154-101-

103), 50°, 219, 29 L, 30 P[UCLA]. Waitangi West, 28 Feb 1964, S (NZ 153),

145 L, 1 P[UCLA]. Locality not specified, Nov 1959, L.J. Dumbleton, 2 0’,

192, 15 L, 2 P [Dumbleton].

AUCKLAND ISLANDS. Tucker Point, 20 Jan 1963, L.J. Dumbleton, 2 L,

1 P, egg raft. Ranui, 28 Dec 1962-22 Jan 1963, L.J. Dumbleton, 50, 2 9,

31 L, 10 P. Ocean Island, 23-29 Dec 1962, L.J. Dumbleton, 1 lpo (L2),

Sip? (Ll, 3,4); 1p? (PA), 3. 1p{(L5-7), 2 p, 2? (14a),.4 o 20,2 2 (14b),

SL, 1 P (14), 3¢%, 29, 4L, 8 P [Dumbleton].

11. Culex (C.) rotoruae Belkin, n. sp.

Figs. 13, 32

Culex (C.) sp. 2, Rotorua form of Belkin (1962:191).

TYPES: Holotype “ (NZ 46-111) with associated larval and pupal skins and

genitalia slide, Kuirau Hot Springs Reserve, Rotorua (South Auckland), New

Zealand, 23-27 Dec 1963, J.N. Belkin and D.A. Schroeder [NELS]. Allotype

? (NZ 46-102) with associated larval and pupal skins, same data as holotype

[NELS]. Paratypes: 26 o, 359, 1251 larvae, 81 pupae (NZ 46), same data as

holotype, as follows: 1 lp? (46-103), 1 o (640124-1), 5 L[CANT]; 5L [NELS];

1 lp? (46-108), 1 o% (640124-2), 5 L [USNM]; 1 1p? (46-110), 1 o% (640124-3),

5 L [BMNH]; 2 1p? (46-117, 119), 14 1p (46-101, 104-107, 109, 112-116, 118, 120,

121), 23¢7, 309, 1226 L, 76 P [UCLA].

FEMALE. Wing: 3.75 mm. Proboscis: 1.75 mm. Forefemur: 1.6 mm.

Abdomen: about 2.6 mm. In general very similar to pervigilans as described

above but distinguished from it by the following features. Size considerably

smaller and general coloration darker. Erect scales of head predominantly

dark, from light brown to black; palpus with very few or no white scales; first

flagellar segment with only a few light scales. Mesonotal integument darker;

mesonotal scaling predominantly dark bronzy not contrasting with integument;

fossa with numerous short curved bronzy scales and without light scales; light

scaling restricted to narrow inconspicuous lines on anterior promontory, lat-

eral prescutal, posterior fossal, supraalar and lateral prescutellar areas; scu-

tellum with light scales as in pervigilans. Pleuron very dark brown; apn and ppn

scales less numerous and darker; ppl, stp, mep scales whitish but less numer-

ous. Tergal abdominal bands narrower, straighter; sternites predominantly

dark-scaled, whitish scales largely restricted to basolateral patches.

MALE. Distinguished from pervigilans by the same characters of thoracic

ornamentation as the female and the following. Palpus essentially as in pervig-

ilans but with long hairs more numerous. Abdomen apparently with more nu-

merous and longer ventral hairs; tergites with larger basal transverse light

markings. Hindfemur with more distinct light line on anterior surface.

Belkin: Culicidae of New Zealand 65

MALE GENITALIA (fig. 13). As figured, apparently indistinguishable from

pervigilans. Leaf of subapical lobe perhaps smaller; 1 or 2 setae in group d-f.

PUPA (fig.13). Abdomen: 2.94mm. Trumpet: 0.6 mm. Paddle: 0.85 mm.

As figured, apparently indistinguishable from pervigilans. Trumpet relatively

short and broad, index usually about 6.0; pinna usually about 0.25 of trumpet

length; uniformly dark or lighter in middle. Hair 5-IV usually shorter than

tergite V. Both paddle hairs usually present.

LARVA (Belkin 1962:fig.92). Head: 0.84 mm. Siphon: 1.26 mm. Saddle:

0.39 mm. As figured by Belkin (1962), in general similar to pervigilans and

resembling quinquefasciatus in the short antenna; distinguished from the for-

mer by the following features. Posterior part of head capsule strongly dark-

ened; hairs 5-7-C shorter and with more branches; mental plate usually with

5,6 teeth on each side. Antenna short and straight, usually less than 0.5 of

head length; uniformly darkly pigmented. Prothoracic hairs 4, 7, 8-P frequent-

ly triple. Abdominal hair 1-III-V weaker; 6-II usually 4, 5b; 6-IJI-VI usually

triple. Comb scales shorter. Siphon lightly pigmented, base and apex dark-

ened; index about 4.0; pecten spines usually with only 2,3 anterior teeth; sub-

ventral tufts (1, la-S) usually 5 pairs (rarely 4, 6,7) all in line on ventral part

of siphon, basal 1, 2 pairs within the pecten, proximal ones usually with about

10 branches which are subequal to width of siphon or shorter. Ventral brush

usually with 7 pairs of hairs, very rarely with 13 hairs; hair 2-X often triple.

Anal gills short, pointed, usually subequal to saddle length or shorter.

SYSTEMATICS. C. rotoruae is most readily differentiated from the other

‘members of the pervigilans complex by the characteristic development of the

Siphon and ventral brush of the larva but it can also be recognized relatively

easily by the dark mesonotal scaling of the adults. There is remarkably little

variation in these diagnostic characters as well as in most other features of the

larva except for the length of the antenna. There is no Significant overlap in

either the adult or the larva with pervigilans or asteliae. On the other hand,

the pupa appears to be indistinguishable from pervigilans and the male genitalia

show no obvious differences from either pervigilans or asteliae.

C. rotoruae is known at present only from Rotorua and immediate vicinity

near the center of the volcanic district which apparently dates from early

Pleistocene. However, the derivation of rotoruae from the pervigilans stock

may have taken place during one of the earlier periods of volcanic activity. It

would be of great interest to determine the entire present range of rotoruae and

any areas of overlap or contact with pervigilans.

BIONOMICS. We found immature stages of rotoruae in very large num-

bers in thermal pools and their outflows with a water temperature ranging from

80° to 83°F (NZ 46). Wallwork collected larvae ina "stagnant pool with a tem-

perature of 80°F" (504) and in ''clear water, temperature 1009F" (503). This

species may breed in cooler water also, since Forsyth found at least one larva

in the stream bounding the Whakarewarewa thermal area in water of "normal

temperature, not thermal" (NZ 185). No other mosquitoes are definitely known

to utilize the same breeding sites as rotoruae; the single rearing of Paradixa

fuscinervis from collection NZ 46 is probably a contamination from a previous

collection.

We had little success rearing adults in the laboratory from our collections

either in the original highly sulphurous water or in fresh tap water. The ma-

jority of larvae died before pupation and few adults emerged even from field-

collected pupae. It appears, therefore, that rotoruae requires Some special

factors for successful growth and metamorphosis which is present only in the

66 | Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

natural breeding sites. There was a remarkable synchrony in the stage of de-

velopment of the immature stages in the field, apparently different in the vari-

ous pools or even parts of pools.

A few adults were found restingin the grass around the breeding sites at the

time our collections were made. Nothing definite is known of the bionomics of

the adults but it seems probable that the 2 females recorded below were taken

landing or biting.

DISTRIBUTION (fig.32). Material examined: 1416 specimens; 27 0, 38 9,

1268 larvae, 83 pupae; 21 individual rearings (7 larval, 14 incomplete).

NEW ZEALAND. South Auckland: Rotorua, Kuirau Hot Springs Reserve,

23,27 Dec 1963, BS (NZ 46) type series as listed above [NELS, CANT, USNM,

BMNH, UCLA]; Whakarewarewa thermal area, 19 May 1964, F (NZ 185), 1 L

[UCLA]. Rotorua, behind baths, hot water, 14 Jan 1944, W.J. Phillips (1),

3 L[NELS]; clear water, temperature 100°F, 3 Jan 1945, Wallwork (503), 6 L

[NELS]; stagnant pool, temperature 80°F, 3 Jan 1945, Wallwork (504), 6 L

[NELS]; 24 July 1923, O.H. Swezey, 19 [BISH]; hot pool, 14 Jan 1944, W.J.

Phillipps, 12 [WELL].

12. Culex (Culex) asteliae Belkin, n. sp.

Figs. 14, 15, 32

TYPES: Holotype “ (NZ 68-104) with associated larval and pupal skins,

Rangitoto Island, 7 Jan 1964, J.N. Belkin and D.A. Schroeder [NELS]. Allo-

type 2 (NZ 68-129) with associated larval and pupal skins, same data as holo-

type [NELS]. Paratypes: 52%, 619, 44 larvae, 122 pupae (NZ 67,68), same

data as holotype except that 4 Jan 1964 for collection NZ 67, as follows: 1 lpo&

(68-102), 1 lp? (68-127) [CANT]; 1 1lpo (68-103), 1 1p? (68-128) [USNM];

1 lpo (68-107), 1 1p? (68-130) [BMNH]; 5 lpo (67-108-110, 114,120), 4 1p?

(67-119, 121-123), 4 po (67-103, 104, 107, 116), 6 p? (67-102, 105, 106, 111, 113,

118), 1 (67), 2:9167); 117 1. (6%), 7 P (67) [UCLA]y 11 1po (68-101, 106; 117,

118, 136, 141, 143, 149, 152, 159, 175), 13 lp? (68-105, 137, 138, 140, 151, 165-168,

174, 178, 181, 182), 24 po (68-111, 115, 119-121, 124, 131, 134, 139, 142, 144-146,

150, 154-158, 160, 162, 164, 169, 177), 24 p? (68-108-110, 112, 113, 116, 122, 123,

125, 126, 131a, 132, 135, 147, 148, 161, 163, 170-173, 176, 179, 180), 1 1p (68-153),

4 (68), 8 2 (68), 297 L (68), 24 P (68) [UCLA].

FEMALE. Wing: 3.5mm. Proboscis: 1.85 mm. Forefemur: 1.6 mm. Ab-

domen: about 2.6 mm. Extremely similar to pervigilans from which it is diffi-

cult to differentiate; distinguished from rotoruae by same features as pervigi-

lans and from the latter primarily by the following. Size considerably smaller

(smallest in complex), general coloration usually lighter. Torus of antenna of-

ten light brown. Mesonotal integument light to dark brown; mesonotal scaling

usually less contrasting, predominantly creamy, whitish toward sides and pre-

scutellar area, supraalar area with a conspicuous patch of bronzy scales, a few

bronzy curved scales in fossa. Forecoxa with a smaller patch of light scales at

base; knee spots and apical hindtibial spot usually yellowish. Abdominal tergal

light bands usually larger and straighter; sternites more extensively dark-

scaled, light scales usually creamy.

MALE. Even more similar to pervigilans than female; mesonotal scaling

essentially same as in pervigilans. Palpus with long hairs shorter and less nu-

merous, those on segment 3 usually less than 10 and restricted to distal third.

MALE GENITALIA (fig. 14). As figured, apparently indistinguishable from

Belkin: Culicidae of New Zealand 67

pervigilans. Seta c of subapical lobe apparently shorter and weaker; 1 to 3

setae in group d-f. Proctiger crown apparently with weaker and fewer spicules.

PUPA (fig. 14). Abdomen: 2.69 mm. Trumpet: 0.58 mm. Paddle: 0.75 mm.

As figured; in general similar to pervigilans and rotoruae from which it can be

readily separated by the following features. Trumpet slender, index at least

8.0; usually uniformly moderately pigmented; pinna less than 0.18. Cephalo-

thorax, trumpet and abdomen pigmented a bright yellowish brown, lighter on >

distal segments. Hairs strongly pigmented, conspicuous; hair 5-IV long, usu-

ally about 1.5 of length of tergite V. Paddle practically colorless except for

strong rib and base; hair 1-P strong, hair 2-P apparently always undeveloped.

LARVA (fig.15). Head: 0.8 mm. Siphon: 1.63 mm. Saddle: 0.42 mm. As

figured; in general similar to pervigilans but easily differentiated by the fol-

lowing features. Head capsule uniformly moderately to lightly pigmented ex-

cept for usual lighter and darker areas; mental plate usually with 8 teeth on

each side, moderately pigmented. Antenna uniformly lightly to moderately

pigmented except for narrow dark basal ring; length usually about 0.63 of head;

spicules shorter and less conspicuous. Prothoracic hairs 7, 8-P usually single,

rarely double. Abdominal hairs 1, 6-III-V usually longer. Siphon moderately

to lightly pigmented, frequently darkened in distal 0.3; long and slender, very

gradually slightly tapered distad, index always more than 8.5; pectenteeth very

small, usually less than 10; subventral tufts (1, 1a-S) usually 4 pairs, the mid-

dle 2 displaced dorsad, the basal and apical more or less in line with pecten,

all tufts usually triple or double and longer than width of siphon. Saddle very

deeply and abruptly incised ventrolaterally for ventral brush; saddle hair (1-S)

usually strong and double. Anal gills subequal, shorter than saddle and pointed.

SYSTEMATICS. C. asteliae is strikingly differentiated from other members

of the pervigilans complex in the larva and pupa but the adults and male geni-

talia are very similar to pervigilans. Particularly distinctive of asteliae is the

long slender larval siphon with the 2 median hair tufts above the level of the

pecten and the basal and apical tufts in line with the pecten. There is remark-

ably little variation in the diagnostic characters of the larva and the trumpet

of the pupa within and among different populations. Therefore, it seems clear

that these characters are genetically fixed and that asteliae is a distinct spe-

cies. I have not seen any larvae of pervigilans which could be mistaken for as-

teliae and there is no indication of any intermediate condition which would sug-

gest hybridization between the 2 forms.

To date asteliae has been collected only in the Hauraki Gulf area and the

adjacent Waitakere Ranges in North Island but its natural range may be coex-

tensive with its principal host plant, the broad-leaf astelia Collospermum has-

tatum. This liliaceous plant on Rangigoto Island is a typical pioneer, growing

on bare volcanic rock on the periphery of the island. I believe that the associ-

ation with this plant may hold the clue to the origin of asteliae and I suggest

that asteliae was formed from a founder population of the pervigilans stock on

an isolated volcanic island where the only available fresh water was in astelia

axils. This event cannot be dated definitely but the relatively wide present

distribution of asteliae suggests that it was not a very recent one, almost cer-

tainly not within the period of man's occupation of New Zealand.

BIONOMICS. The immature stages of asteliae have been collected only in

the leaf axils of terrestrial and epiphytic broad-leaf astelias, principally

Collospermum hastatum but possibly also one or more species of the genus

Astelia. We did not find asteliae in other plants which contain axil water in

smaller quantity and less regularly, such as nikau palms, Phormium, Cordy-

68 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

line and narrow-leaf species of Astelia but it is possible that it may breed

in these occasionally. The only other mosquito definitely known to breed in

leaf axils of astelias is Aedes notoscriptus which we found once in association

with asteliae (NZ 67) and which has been reported before from astelias (Miller

1920:36; Graham 1929:210). The association of quinquefasciatus with asteliae

in one collection (NZ 93) is probably due to contamination or error in labeling.

The immature stages of asteliae were reared without difficulty in the

laboratory but this species appeared to require more time than pervigilans

for the completion of the aquatic cycle. Nothing is known of adult bionomics

Since this species has never been collected in the field in this stage, even on

Rongitoto Island where the immature stages are quite common.

DISTRIBUTION (fig.32). Material examined: 850 specimens; 67 0, 69 9,

909 larvae, 159 pupae; 125 individual rearings (58 larval, 63 pupal, 4 incom-

plete).

NEW ZEALAND. North Auckland: Rangitoto Island, 14 Sept 1963, S (NZ

7), 11lpo (7-112), 4 L[UCLA]; 4 Jan 1964, S (NZ 67), 7 Jan 1964, BS (NZ 68),

type series as listed above [NELS, CANT, USNM, BMNH, UCLA]. Waitakere

Ranges, Karekare Road at crossing of headwaters of Opal Pools, 8 Jan 1964,

BES (NZ 82), 1 lpo& (82-102), 1 L, 1 P[UCLA]. Titirangi Beach, end of Ma-

hoe Road, 11 Jan 1964, BS (NZ 93), 1 lpo& (93-101), 1 1p? (93-102), 1 1p (93-

104), 4L, 2 P [UCLA].

Little Barrier Island: Summit Track, elev. 186 m, 10 Oct 1963, S (NZ 13),

3 lpo (13-103, 105, 106), 1 po (13-101), 1 1p (13-107), 8 L: [UCLA]; ‘8 Nov

1963, S(NZ 25), 3 lpo (25-101, 106, 110), 1 lp (25-102), 8 L[UCLA]. Summit

Track, elev. 267m, 8 Nov 1963, S (NZ 24), 3 lpo& (24-101, 102, 104), 1 1p?

(24-103), 9 L[UCLA]. Summit Track, elev. 75m, 8 Nov 1963, S(NZ 26), 1 9,

3 L[UCLA]. Summit Track, elev. 225m, 7 Mar 1964, S(NZ 132), 1lpo

(132-103), 1 1p? (132-107), 1 po" (132-106), 3 p? (132-102, 104,105), 41 L,1P

[UCLA ].

South Auckland: Coromandel, 22 Apr 1964, N (NZ 219), 2 L [UCLA].

13. Culex (Culex) quinquefasciatus Say

Fig. 33

1823. Culex 5-fasciatus Say, 1823:10. TYPE: Sex not specified [New Orleans,

Louisiana], Mississippi River, U.S.A. [Lost]. Synonymy with fatigans

not universally accepted, see Stone (1957:342-343); type locality re-

stricted by Belkin, Schick and Heinemann (1966:4-5).

1828. Culex fatigans Wiedemann, 1828:10. TYPE: Lectotype ? bearing Wiede-

mann's handwritten label (lacking on o which also lacks the genitalia), as

follows/Ind. orient/Coll. Winthem/fatigans Wied/Ind orient/fatigans/

det. Wiedem. //; PRESENT SELECTION [NMW]. Synonymy with quinque-

fasciatus by Dyar and Knab (1909:34), not universally accepted, see Stone

(1957:342-343).

1848. Culex acer Walker, 1848:18. TYPE: Holotype 2, New Zealand [BMNH].

Synonymy with fatigans by Edwards (1924:336, 395).

For extensive complete synonymy, see Stone, Knight and Starcke (1959).

C. (C.) quinquefasciatus of Belkin (1962:195-197); Stone (1963:135).

C. (C.) pipiens quinquefasciatus of Stone, Knight and Starcke (1959); Stone

(1967:217).

C. (C.) fatigans of Edwards (1924:395; 1928:47; 1932:208); Taylor (1934:24);

Halkins Culicidae of New Zealand 69

Graham (1939:211); Miller and Phillipps (1952:21).

C. (C.) pipiens fatigans of Laird (1956).

FEMALE (Miller and Phillipps 1952:fig. 37,38). Wing: 3.9 mm. Probos-

cis: 2.15 mm. Forefemur: 2.0 mm. Abdomen: about 2.85 mm. As described

by Beikin (1962:195-196); in general similar to members of pervigilans com-

plex; differing from pervigilans chiefly in the following diagnostic features.

General coloration lighter, integument and dark scales lighter, light scales

usually creamy instead of white. Thoracic integument lighter, mesonotum uni-

formly brown, pleuron light brown; mesonotal scales shorter and more numer-

ous, darker in color, largely light brown and not contrasting with integument.

Anterior surface of hindfemur entirely pale-scaled at base and on lower surface

distally to near apex; posterior and ventral surfaces of all femora largely pale-

scaled; knee spots and hindtibial apical spot less conspicuous and more light

tan than white; lower surfaces of tibiae and sometimes basal tarsal segments

more or less distinctly paler than upper surfaces. Abdominal tergal light scales

usually creamy rather than whitish, basal transverse light bands usually more

strongly widened in the middle; sternites I-VII usually with pale tan scales on-

ly, rarely with a few dark scales in the middle of some segments as in the male

(possibly hybrids with pervigilans).

MALE, Essentially as in the female but generally paler. Differing from

pervigilans in the same features as the female except for the following. Ab-

dominal sternites II-VII with some dark scales in the middle usually in the

basal 0.5 only, apical half of each segment pale-scaled straight across, with-

out dark scales in posterolateral areas. Palpus apparently somewhat shorter,

rarely exceeding proboscis by more than length of segment 5; distal segments

with less numerous and shorter hairs; segment 3 usually with less than 15

moderately long external ventrolateral hairs in distal half or less, a more or

less distinct pale-scaled line usually present above the hairs.

MALE GENITALIA (Belkin 1962:fig.97). As figured and described by

Belkin (1962:196) and with the following diagnostic features from the pervig-

ilans complex. Tergal arms of the outer division of lateral plate of the phallo-

some relatively short and straight, subparallel, apex acuminate in dorsal as-

pect, DV/V usually more than 0.7. Basal sternal process short but variable,

usually poorly pigmented, rarely moderately pigmented and somewhat elon-

gate but never strongly curved ventrad.

PUPA (Belkin 1962:fig.97). Abdomen: 3.27 mm. Trumpet: 0.8 mm.

Paddle: 0.9mm. As described by Belkin (1962:196); extremely similar to

members of pervigilans complex from which it can be distinguished apparently

by the strongly oblique pinna whose maximum length is at least 0.4 of the total

length of the trumpet.

LARVA (Miller and Phillipps 1952:fig.39; Belkin 1962:fig.98). Head:

0.79 mm. Siphon: 1.02 mm. Anal Saddle: 0.4 mm. As figured’and described

by Belkin (1962:196); in general similar to pervigilans from which it is readily

distinguished by the following characters. Mental plate usually with at least

9 teeth on each side of median apical tooth; aulaeum much shorter and less

conspicuous; head capsule usually moderately to strongly uniformly pigmen-

ted. Antenna much shorter (as in rotoruae), usually 0.6 or less of head cap-

sule; moderately to strongly pigmented, middle of proximal part lighter, base

darkened; spicules well developed dorsally on proximal part, slender except

toward hair 1-A, distal part usually without spicules. Abdominal hair 1

usually single or double on III-V. Siphon more or less distinctly swollen

70 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

near or before middle, apex narrowed; shorter, index usually less than 5.0;

subventral tufts (1, la-S) usually 4 pairs, subapical distinctly out of line, prox-

imal distad of pecten.

SYSTEMATICS. The ubiquitous pipiens complex is apparently represented

in New Zealand only by small populations of quinquefasciatus which were

undoubtedly introduced in ships. In most features these populations are quite

Similar to populations of quinquefasciatus in the tropical South Pacific and else-

where in the world. However, in one lot of larvae (NZ 69), there is a sugges-

tion of introgression of pervigilans characters, particularly in the shape and

number of teeth in the mental plate. Although these characters are some-

what variable in extralimital populations and may be temperature dependent,

the possibility of hybridization of quinquefasciatus and pervigilans should not

be dismissed without experimental studies.

It is possible that other forms of the pipiens complex exist in New Zealand.

The females from Waitomo are undoubtedly members of this complex but their

assignment to quinquefasciatus is provisional only since all forms in the com-

plex are recognized almost exclusively on male genitalic characters.

BIONOMICS. C. quinquefasciatus appears to be uncommon in New Zealand

and largely confined to coastal areas in the vicinity of the larger ports on North

Island where it appears to be definitely established. Our collections of imma-

ture stages were made in breeding sites typical of this form in other parts of

the world, ina fire drum (NZ 66), a cistern cover and air hole (NZ 69), an ox-

idation pond in a Sewage treatment plant (NZ 180), an abandoned washing ma-

chine bowl (NZ 194), and a boat hull (NZ 218). The pupa reported from an as-

telia axil (NZ 93) is undoubtedly quinquefasciatus but this record may be due to

contamination or an error in labeling. On only one occasion (NZ 66) was quin-

quefasciatus collected in association with pervigilans but the 2 species may oc-

cur together more frequently since there is a suggestion of introgression of

pervigilans characters in several specimens in another collection of quinquefas-

ciatus (NZ 69).

Our collections do not support Graham's (1939:211) statement that this spe-

cies breeds ‘'through the year in any place exposed to full sunlight and holding

water charged with organic matter.'' Graham (loc. cit.) states that he found

quinquefasciatus (as fatigans) in the Auckland District breeding and biting all

through the winter and observed pupation and emergence of adults in June.

Adults, however, were not encountered in September and October, and he be-

lieved that the months of July and August were cold enough to kill them and

suggested that the species survived from year to year through "hibernation" of

larvae.

DISTRIBUTION (fig. 33). Material examined: 244 specimens; 40 c, 43 9,

117 larvae, 44 pupae; 18 individual rearings (16 larval, 1 pupal, 1 incomplete).

Widespread in tropical, subtropical and warm temperate regions.

NEW ZEALAND. North Auckland: Whangarei, Feb 1924, T.R. Harris,

12[BMNH]. Devonport, 18 Feb 1917, D. Miller, 1 o [BMNH]; Shoal Bay, 18

Jan 1917, D. Miller, 1c [BMNH]. Rangitoto Island, 7 Jan 1964, BS (NZ 69),

5 ipo (869-102-106), 3 lp? (69-101, 107, 108), 2c, 11°, 66, 14 P [UCLA],

Oratia, DSIR Field Station, 30 Dec 1963, S (NZ 66), 1 1p (66-103), 18 L[UCLA].

Titirangi Beach, end of Mahoe Road, 11 Jan 1964, BS (NZ 93), 1 P [UCLA].

Auckland, Avondale, Methuen Road, 9 Aug 1964, F (NZ 194), 1 po (194-124)

[UCLA]. Glendion(?), "in banana palm,'' 20 Mar 1919, D. Miller, 1 “|BMNH].

Erin Bay(?), 24 Mar 1919, D. Miller, 1c [BMNH]; Erin Park(?), same data,

"catch basin, '' 1 o [BMNH]. Mangere Sewage Plant, 22 Apr 1964, F (NZ 180),

Belkin: Culicidae of New Zealand ts

4 1po" (180-101, 103, 107, 108), 4 1p (180-102, 104, 105, 109), 23 oc, 239, 8 L,

20 P [UCLA].

South Auckland: Coromandel, 21 Apr 1964, N (NZ 218), 8 L[UCLA]. Wai-

tomo, 26 July 1923, G.A. Marshall, 1 ? [BMNH].

TRIBE CULISETINI

Genus CULISETA

1902. Theobaldia Neveu-Lemaire, 1902:1331-1332. TYPE SPECIES: Culex an-

nulatus Schrank, 1776, Europe; original designation. Preoccupied by

Theobaldia Fischer, 1885; invalid emendation for Theobaldius Nevill,

1878 but available name for homonymy.

1904. Culiseta Felt, 1904:391c. TYPE SPECIES: *Culex absobrinus Felt, 1904,

New York; original designation.

1915. Climacura Howard, Dyar & Knab, 1915:452. TYPE SPECIES: *Culex

melanurus Coquillet, 1902, New Hampshire.

For complete synonymy see Stone, Knight and Starcke (1959).

14. Culiseta (Climacura) tonnoiri (Edwards)

Fig. 34

1925. Theobaldia tonnoiri Edwards, 1925:258-259. TYPE: *Holotype 2, Waiho,

Westland, N. Z., 16 Jan 1922, A.L. Tonnoir [NELS]. |

Culiseta (Climacura) tonnoiri of Belkin (1962:284-286); Stone (1961:44); Nye and

McGregor (1964:7-8); Dumbleton (1965:137-142); Pillai (1966).

Culiseta (Culiseta) tonnoiri of Stone, Knight and Starcke (1959:220).

Theobaldia (Climacura) tonnoiri of Edwards (1932:106); Taylor 1934:13).

Theobaldinella tonnoiri of Miller (1950:45); Miller and Phillipps (1952:25-26);

Ross, Austin et al (1963:20).

FEMALE (Miller and Phillipps 1952:fig. 44-46). Wing: 4.7mm. Proboscis:

3.15 mm. Forefemur: 2.25 mm. Abdomen: about 3.0 mm. As described by

Belkin (1962:284-285), with the following diagnostic features. Scales of vertex

of head all golden; labium and palpus dark-scaled. Mesonotal, scutellar and

pleural integument dull reddish brown, pleuron with lighter areas; mesonotal

scales all linear, largely golden but dark brown in large patches on fossa, pos-

terior dorsocentral and upper supraalar areas; pleural scaling very scanty,

narrow hairlike golden scales on apn and ppn, elongate broad semierect creamy

scales on upper and lower patch on stp and patch in middle of meép; 3 to 5 spi-

racular bristles, 1 strong lower mep bristle. Legs dark brown; femora with a

subapical light ring; ventral and posterior surfaces of forefemur and midfe-

mur and all but dorsal surface of hindfemur light-scaled. Claws all simple and

all with long fine spicules on outer surface. Wing with distinct large dark

spots around furcation of Rs and adjacent veins, on furcation of Rg+3 and fur-

cation of M, formed by numerous scales which are darker than elsewhere;

membrane of wing conspicuously infuscated in area of Rs spot; vein Sc witha

large patch of long bronzy bristles at base on lower surface. Abdomen with dark

scales only but appearing banded on base of segments because of sparser scaling.

72 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

MALE (Dumbleton 1965:figs. 1,2). As described by Dumbleton (1965:138);

in general similar to female except for sexual characters. Palpus as long as

proboscis or slightly shorter, with dark scales on segments 2-4, segments 4

and 9 subequal. Claws of forelegs and midlegs enlarged, unequal, larger of

pair with basal and submedian teeth.

MALE GENITALIA (Dumbleton 1965:fig.3). As described by Dumbleton

(1965:138). Ninth tergite lobe with 3, 4 long, heavy bristles. Sidepiece without

scales; basal mesal lobe poorly differentiated, not projecting, with specialized

bristles. Clasper about as long as sidepiece, very slender and slightly tapering.

Aedeagus large, about twice as long as wide, simple except for acute apex pro-

jecting tergally. Paraproct with 2 conspicuous stout curved teeth; cercal setae

PUPA (Dumbleton 1965:figs. 4-6). As described by Dumbleton (1965:140)

_ from a single damaged skin. The following combination of generic and subge-

neric characters evident in this skin will separate the species from all other

known New Zealand Culicinae. Trumpet with a long slit in the meatus, hair 9-

Vill at the posterolateral angle of the segment, hair 1-IX a simple hair, both

paddle hairs (1,2-P) present. The external and internal margins of the paddle

have distinct denticles.

LARVA (Belkin 1962:fig.185; Miller and Phillipps 1952:fig.47; Nye and

McGregor 1964:fig. 5; Dumbleton 1965:fig. 7). Head: 0.95 mm. Siphon: 1.96

mm. Anal Saddle: 0.51 mm. As figured and described by Belkin (1962:285);

with the following diagnostic features. Head lightly to moderately pigmented;

mental plate short and broad, with 9,10 teeth on each side; hair 6-C single. An-

tenna uniformly darkly pigmented except for black basal ring; longer than head

capsule, slender, uniform in width to hair 1-A, narrowed beyond and again dis-

tad of hairs 2,3-A. Thorax said to be whitish in life and abdomen reddish or

purplish (Miller and Phillipps 1952:26). Comb with about 18-21 scales (possibly

up to 27 or more) arranged in a single regular row, median scales with black

central rod and transparent apex and sides. Siphon variable in length, index

6.5-9.0; uniformly lightly to strongly pigmented, with darkened apex and black-

ish base and acus; pecten teeth usually 8-10, simple or rarely with 1, 2 minute

subbasal denticles; hair 1-S very small, at extreme base of siphon, usually

3-5b; hairs la-S in a single midventral row, very fine, short, single or double;

hair 2-S a rather strong slender apically hooked stiff hair at extreme apex of

siphon body; hairs 2a-S in an irregular subdorsal row, 6-8 on each side, very

fine, short, single or double. Saddle complete; length and width apparently

variable, possibly a sexual character; saddle hair (1-X) small, fine, usually

3b; ventral brush with 7 pairs of hairs distad of saddle, 6 pairs of these on

grid, usually 3,4 short hairs on saddle, mostly thick, single or double. Anal

gills apparently variable, simple or distinctly constricted near middle.

SYSTEMATICS. Although females of Culiseta tonnoiri are very common

in West Otago and Westland, the male and theimmature stages are poorly known.

I have seen a single male from an individual larval rearing (described by Dum-

bleton 1965). Although the possibility exists that more than 1 species of Cli-

macura is present in New Zealand, I believe that the differences in the larvae

(constriction of gills, number of comb scales, character of pecten teeth, length

and proportions of the siphon and anal saddle) are probably individual or sex

dependent variations in a single species. My illustration (Belkin 1962:fig. 185)

was made from a strongly flattened larva and the siphon index, the proportions

of the anal saddle and the anal gills are distorted. The 2 larvae from South

Westland collected by Dumbleton (1965:140) agree very well with the material

Belkin: Culicidae of New Zealand 73

from Waipoua Forest except for the terminal segments and even in these 2 lar-

vae there is considerable variation in the number of branches in the hairs of

abdominal segment VIII. Therefore, I believe that only one highly variable

species is present in New Zealand. However, special efforts should be made

to rear individually adequate samples from different populations of Culiseta to

determine the range of variation in tonnoiri.

Culiseta tonnoiri is unquestionally related to antipodea Dobrotworsky, 1962

from Southeast Australia and to melanura Coquillett, 1902 from the Eastern and

Central United States, the only other species of Culiseta which can be referred

to the subgenus Culiseta at this time. The correspondence between melanura

and tonnoiri in all the significant characters of the larva and pupa is truly re-

markable and can hardly be due to convergence. Dumbleton (1965:137) has

pointed out that a similar distribution pattern is known for amphibians and plants.

BIONOMICS. Dumbleton (1965:141-142) discusses the habitat, breeding

places, biology and the possible relation of Culiseta tonnoiri to disease trans-

mission in man and animals. On North Island tonnoiri larvae have been col-

lected only once on 19 March 1945 from "'back waters of streams where there

is very slow-flowing, or comparatively dead, water sheltered by a dense cano-

py of foliage and rich in decaying matter including leaves and other debris;

among the latter, the larvae of tonnoiri, like those of antipodeus, hide when

alarmed and are difficult to find, but unlike antipodeus they appear to develop

chiefly during the summer" (Miller and Phillipps 1952:26). In Westland, 2 lar-

vae were collected in January and February 1965 in "permanent ground pools

under open forest. The pools appeared to be black, but the water was actually

a light amber colour" (Dumbleton 1965:141). Dumbleton suggests that tonnoiri

is univoltine and breeds primarily in November and December and hibernates

as a fertilized female.

This species is found primarily in areas of native forest. Females have |

been taken landing and biting in December, January and February in Westland

and in February and June on North Island. Dumbleton (loc. cit.) reports a few

females taken in light traps and in buildings in Westland in October. Biting

activity starts shortly after dark. The natural hosts of tonnoiri are probably

birds. Ross, Austin, et al (1963) have isolated viral agents from tonnoiri.

DISTRIBUTION (fig. 34). Material examined: 90 specimens; 1, 59 9, 29

larvae, 1 pupa; 1 individual larval rearing.

NEW ZEALAND. North Auckland: Waipoua Forest, H.R. Simpson, 19 Mar

1945 (642), 2 L[UCLA]; 19 Mar 1945 (657), 4 L [NELS]; June 1945 (744), 2 L

[UCLA], 19, 9 L[NELS]; June 1945 (745), 19 2 [NELS]; June 1945 (746), 10 L

[NELS]. Auckland Watershed Reservoir, 26 Feb 1944, G. Chamberlain, 1 9

[WELL].

Nelson: Maruia Springs, 22 Jan 1958, L.J. Dumbleton, ?[Dumbleton 1965].

Westland: Lake Moana (Brunner), 16-21 Dec 1925, A.L. Tonnoir, 6 9

[CANT]; Lake Ianthe, 11 Jan 1965, L.J. Dumbleton, ? [Dumbleton 1965]. Lake

Mapourika, 7 Feb 1964, S(NZ 112), 279 [UCLA]; 1 Feb 1965, L. J. Dumble-

ton, ¢ [Dumbleton 1965]. Waiho, 16 Jan 1922, A.L. Tonnoir, ? holotype, 1 ?

[NELS], 19[BMNH]. Franz Josef Glacier, 30 Oct 1963, L.J. Dumbleton, ?

[Dumbleton 1965]. Lake Matheson, 16 Dec 1963, L.J. Dumbleton, ? [Dumble-

ton 1965]. Karangarua River, Mar 1950, L.J. Dumbleton, 2 [Dumbleton 1965].

Copland River, Welcome Flat, Mar 1950, L.J. Dumbleton, ? [Dumbleton 1965].

Paringa River, 18 Nov 1950, L.J. Dumbleton, ? [Dumbleton 1965]. Blue River,

Lake Moeraki, 12 Jan 1965, L.J. Dumbleton, 1 L; 3 Feb 1965, L.J. Dumble-

ton, 11po [Dumbleton 1965]. Haast, 27 Jan 1962, L.J. Dumbleton, ? [Dumble-

714 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

ton 1965]. |

Otago: Martin's Bay, 1-4 Feb 1955, R.R. Forster, 2? [AUCK]. South

Otago, locality not specified, L[J.S. Pillai, in litt. May 1965].

Southland: Milford Sound, Dec 1945, L.J. Dumbleton, ? [Dumbleton 1965].

TRIBE MANSONINI

Genus COQUILLETTIDIA

1905. Coquillettidia Dyar, 1905:47. TYPE SPECIES: *Culex perturbans Walker,

1856 [Charleston, South Carolina], U.S.A. (restriction by Belkin, Schick

and Heinemann 1966:3); original designation.

FEMALES. As described by Belkin (1962:303) and with the following dis-

tinctive features in New Zealand species. Labium entirely dark-scaled; palpus

usually entirely dark-scaled, sometimes with a few pale scales; apical palpal

segment minute, without scales. Acrostichal bristles in a complete row; psp

bristles absent. Tarsi without light rings. Dorsal scales of wing veins all dark

and narrow. Abdominal tergites largely dark-scaled but with basal lateral

patches of light scales.

FEMALE GENITALIA. As described by Belkin (1962:303) except that cer-

cus may be upturned and postgenital plate bilobed (tenuipalpis).

MALES. In general as in the females. Palpus about as long as proboscis

or distinctly longer, segment 5 well developed.

MALE GENITALIA. As described by Belkin (1962:303) and with the fol-

lowing distinctive features in New Zealand species. Sidepiece simple, more or

less cylindrical or conical, without lobes or line of thickened bristles. Clasp-

ette short, poorly developed, bearing 1 or more elongate apical spiniforms.

Clasper irregular, with 1 apical or subapical spiniform. Aedeagus with spic-

ules or denticles. |

PUPAE. As described by Belkin (1962:303) and with the following distinc-

tive features in New Zealand species. Trumpet on a slight tubercle; apex pro-

duced into a poorly sclerotized narrow process which is surrounded by a ''cel-

lular'' lobe on 2 sides, this portion usually broken off in detached pupae. All

hairs short and inconspicuous, usually all unbranched including float hair (1-I).

All hairs (1-12-C) present on cephalothorax, all normal culicine hairs present

on abdomen except 3-VII or 2, 3-VII; hair 1-IX present in iracunda. Paddle

without hairs; inner or both inner and outer margins produced beyond apex of

midrib. |

LARVAE. Only first-instar larvae known for New Zealand species and

these are similar to extralimital species which are readily distinguished from

all other culicid larvae by characters indicated by Belkin (1962:303) and espe-

cially by the unique development of the siphon as follows: short, triangular in

outline, without pecten; ventrolateral valve greatly developed, at least as long

as body of siphon, with recurved spines near and at apex; median valve witha

long sawlike process.

SYSTEMATICS. I am following Ronderos and Bachmann (1963) in recog-

nizing Coquillettidia as a distinct genus instead of retaining it as a subgenus of

Mansonia. It may be even advisable in the future to split Coquillettidia into 2

or more genera but this must await better knowledge of the immature stages

Belkin: Culicidae of New Zealand 15

which are still unknown for most Species.

In New Zealand, Coquillettidia is represented by 2 very distinct endemic

Species, one of which, tenuipalpis, is so different from typical Coquillettidia

in numerous basic characters that it is recognized here as constituting a new

monotypic subgenus (see below). The other species, iracunda, appears to be

closely related to the Australian linealis (Skuse, 1889), and these 2 species

share many characters with perturbans, the type species of Coquillettidia, but

are very different from the majority of palaeotropic species of the crassipes

group. The latter group should probably be segregated into another subgenus

of Coquillettidia.

The 2 New Zealand species have very wide, nearly co-extensive distribu-

tions on North Island and South Island but neither shows any geographical dif-

ferentiation.

BIONOMICS. Both New Zealand species of Coquillettidia breed in the

shallow margins of ponds and small lakes. To date their larvae have not been

collected in nature and only the first instar larvae have been obtained from egg

rafts by Dumbleton (1967, in litt.).

Females attack man readily in late afternoon and at night in the vicinity of

breeding sites where both sexes may be found in large numbers resting in the

daytime on herbs, grasses and brushy vegetation.

Keys to Species

Adults

T: Base of vein Sc below with large group of bristles; apn and ppn without

distinct scales but with a few short hairs (Austranmnsonal :

eer ae tenuipalpis

Base of vein Sc below with scales only: apn and ppn with narrow scales

(COU ea OG ee ee ae nape

Male Genitalia

Ly Aedeagus long and narrow, nearly 5.0 of maximum subapical width;

clasper without outer lobe (Austromansonia). . . . . .1. tenuipalpis

Aedeagus short and broadened subapically, about 2.0 of maximum sub-

apical width; clasper with an outer lobe in addition to small dorsal

one. (Coguilleuigia) a. cise secre Ae co a oe as

Pupae

igs Apex of paddle produced only on the inner margin; hair 9-VIII at apex of

caudolateral angle, or distad; hair 1-IX vagy absent (Austro-

mansonia) . : . . .1. tenuipalpis

Apex of paddle ptoduced beyond mdeib on ate inner and outer margins;

hair 9-VIII distinctly eee Be of apex of caudolateral angle; hair 1-

IX distinct (Coquillettidia) . cae fo ee sacurnda:

Larvae

Unknown

76 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

Austromansonia Belkin, n. subgenus

TYPE SPECIES: Taeniorhynchus (Coquillettidia) tenuipalpis Edwards, 1924,

New Zealand. Only included species.

Austromansonia differs from all other Coquillettidia in the following strik-

ing features: in the adults, vein Sc below with a large patch of hairs as in Culi-

seta, Opifex and Aedes (Nothoskusea), stp and base of pra with a continuous

patch of scales, all claws with long fine Spicules on outer surface of base; in

the male, palpal segments 4 and 5 slightly upturned and with sparse bristles,

flagellar segments 12 and 13 together distinctly longer than remainder of flagel-

lum; in the male genitalia, sidepiece without indication of mesal membrane or

lobes, aedeagus long; in the pupa, only the inner margin of the paddle produced

at apex, hair 9-VIII at apex of caudolateral angle, hair 2-VII present.

Austromansonia Seems to be the most primitive and generalized phylad of

Coquillettidia and of the entire tribe Mansoniini. The pupal stage has the most

complete chaetotaxy of the entire tribe, lacking only hair 3-VII (which is fre-

quently represented by an alveolus) and apparently hair 1-IX. The only other

phylad with nearly as complete a pupal chaetotaxy is the iracunda group of Co-

quillettidia represented by the nominate species in New Zealand and linealis

(Skuse, 1889). in Australia. This group, however, is markedly different from

Austromansonia in general adult morphology and male genitalia as indicated

under iracunda below.

15. Coquillettidia (Austromansonia) tenuipalpis (Edwards)

Figs. 16,17, 35

1924. Taeniorhynchus (Coquillettidia) tenuipalpis Edwards, 1924:336-367.

TYPE: *Holotype © with attached genitalia mount, Ohakune, New Zealand,

Jan 1924, T.R. Harris [| BMNH].

Coquillettidia (C.) tenuipalpis of Ronderos and Bachmann (1963:50); Stone

(1967).

Mansonia (Coquillettidia) tenuipalpis of Edwards (1932:119); Taylor (1934:14);

Stone, Knight and Starcke (1959); Belkin (1962:305); Nye and McGregor (1964:

8-11).

Mansonia tenuipalpis of Miller (1950:45); Miller and Phillipps (1952:24).

Taeniorhynchus tenuipalpis of Graham (1939:212).

FEMALE (fig.16). Wing: 4.83 mm. Proboscis: 3.0 mm. Forefemur:

2.33 mm. Abdomen: about 2.67 mm. As figured here and as described by

Belkin (1962:305); with the following diagnostic features. Narrow decumbent

scales of vertex pale beige, densely packed along eye margins; broad scales

similarly colored, rather elongate; erect scales somewhat darker, rather short.

Mesonotal integument dark brown; mesonotal and scutellar scales all golden;

pleural scales dingy translucent whitish or pale beige. Abdominal tergites

largely bronzy dark-scaled and with extensive basal lateral patches of light

scales; sternites largely light-scaled throughout; sternite I completely unscle-

rotized; sternite II less than 0.5 of III.

FEMALE GENITALIA (fig. 16). Terminal segments very short and deeply

retracted into segment VIII. Tergite VIII about 0.5 of VII, with numerous

heavy bristles caudolaterally. Sternite VIII subequal to VII and about 2.0 length

of tergite VIII, with numerous smaller bristles along caudal margin and in the

middle. Membrane between segments VIII and IX very long, particularly be-

Belkin: Culicidae of New Zealand 17

tween tergites, and containing distinct very slender sclerotized bars (possibly

representing intersegmental lines). Insula and sigma represented by a very

broad plate articulating with cowl; insula with many short bristles. Tergite IX

very narrow, about 0.16 of tergite VIII at its broadest, without bristles in mid-

dle, lateral areas bearing 8-12 bristles on each side in tergal half. Cowl with

longitudinal basal part articulating with tergite IX and sigma, distal partarched.

Tergite X not sclerotized. Cercus compressed, unsclerotized on most of

mesal surface; upturned, thumblike, rounded or slightly truncate apically, and

bearing many short bristles. Postgenital plate separated by membrane from

cowl; small, with 2 distinct lateral lobes bearing numerous short bristles.

Atrial plates absent.

MALE (fig.16). As figured here and as described by Belkin (1962:304-

305); essentially as in the female and with the following distinctive features.

Palpus slightly upturned, about 1.25-1.3 of proboscis, exceeding latter by full

length of segment 5 which is slightly longer than segment 4; apex of segment 3,

all of 4 and base of 5 with a few very short hairs. Antenna slightly shorter than

proboscis; flagellar segments 12 and 13 together about 1.2 of remainder of

flagellum, segment 13 about 1.5 of 12. Claws as figured.

MALE GENITALIA (Belkin 1962:fig. 196; Nye and McGregor 1964:fig. 6).

As figured and described by Belkin (1962), with the following corrections and

diagnostic features. Segment IX: Tergite lobe poorly defined, with 5-8 bristles;

sternite very broad, widely separating sidepieces, with 1 to several pairs of

bristles. Sidepiece simple, without any indication of lobe and mesal mem-

brane. Claspette with very long sternal process projecting between sidepieces

to nearly base of aedeagus; apex with 2 long spiniforms, one of which is con-

siderably thinner than other. Clasper with a conspicuous postmedian and small

preapical dorsal lobe but without outer lobe; base without dorsal ledge. Aedea-

gus long, cylindrical, poorly sclerotized on dorsal surface; a subapical dorsal

process with about 16 short sharp spicules directed cephalad. Paraproct with

about 3-6 spines; cercal setae about 5.

PUPA (fig.17). Abdomen: 4.54 mm. Trumpet: 0.84 mm (without decid-

uous apex). Paddle: 1.34 mm. Chaetotaxy as figured; diagnostic characters

as in the key; all normal culicine hairs present on cephalothorax and abdomen

except 1-IX and 3-VII (frequently represented by an alveolus). Cephalothorax

lightly pigmented except for darkened dorsal surface; metanotum darkly pig-

mented. Trumpet very darkly pigmented on tracheoid; bulb (meatus beyond

tracheoid to base of triangular sclerite) moderately pigmented; neck (from tri-

angular sclerite to apex of non-deciduous part) not markedly narrowed, lightly

pigmented and with smooth apical margin; deciduous apex lacking in all speci-

mens. Abdomen uniformly darkly pigmented; hair 9-VIII at or caudad of prom-

inent caudolateral angle. Paddle uniformly moderately or darkly pigmented;

inner margin strongly produced beyond outer.

LARVA. Unknown.

SYSTEMATICS. As indicated under the diagnosis of Austromansonia, tenui-

palpis possesses a number of striking features in all known stages which set it

apart from all other members of the genus Coquillettidia. In most of these fea-

tures it appears to be the most primitive living representative of the tribe Man-

soniini. Should its larva prove to be as distinctive as the other stages it might

be advisable to raise Austromansonia to full generic rank.

BIONOMICS. Pupal exuviae of tenuipalpis have been found in association

with those of iracunda in shallow margins of ponds and small lakes choked with

herbaceous and grassy vegetation, particularly sedges and reeds. No larvae

were found in these areas in spite of repeated search at different times of the

year.

78 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

Both sexes were swept in numbers from the vegetation in the breeding sites

and adjacent banks. A few females have been collected attempting to bite in the

vicinity of breeding sites and 3 females were caught in a light trap.

This species does not seem to be as abundant as iracunda and apparently

seldom bites man.

DISTRIBUTION (fig.35). Material examined: 176 specimens, 69 o, 97 9,

10 pupal skins; no individual rearings.

NEW ZEALAND. North Auckland: Waimauku, Jan, egg laying, D.H. Gra-

ham (Graham 1939:212). Bethells Road Swamp, 28, 29 Dec 1963, BFS (NZ57A,

97B), 350°, 579, 1 p[UCLA]; 10 Jan 1965, F (NZ 200A), 1c, 49 [UCLA]; 6

Dec 1966, L.J. Dumbleton (NZ 225), 9 p[UCLA]. Waitakeres, 2 Feb 1950,

M. Knight, 19 [AUCK]; 12 Feb 1950, 19[UCLA]; 18 Feb 1950, 1 9 [AUCK],

19[UCLA]; 25 Feb 1950, 22 [AUCK]; 26 Feb 1950, 19[AUCK]. Piha, May,

egg laying, D.H. Graham (Graham 1939:212). Titirangi, 4,14, 18 Feb 1955,

light trap, C.R. Thomas, 3 2 [AUCK]. :

Wellington: Ohakune, elev. 2060 ft, 10 Jan 1920, T.R. Harris, 20, 19

[BMNH]; 15 Dec 1922-15 Jan 1923, T.R. Harris, 19 [BMNH]; Jan 1924, T.R.

Harris, holotype %, 20, 42[BMNH], 1c, 12[UCLA]; pond on bend of Lake

Road, 9-10 Jan 1964, BFS (NZ 85A), 27%, 16 2 [UCLA].

Otago: Martins Bay, 1-4 Feb 1955, R.R. Forster, 1 9 [UCLA].

Southland: Te Anau, in car, 2 Feb 1964, 19 [UCLA].

Subgenus Coquillettidia

16. Coquillettidia (C.) iracunda (Walker)

Figs.17, 35

1848. Culex iracundus Walker 1848:6. TYPE: *Holotype ? (42-55), New Zealand;

identified as the type by E.A. Waterhouse [BMNH].

Coquillettidia (C.) iracunda of Ronderos and Bachmann (1963:50); Stone (1967).

Mansonia (Coquillettidia) iracunda of Edwards (1932:118); Taylor (1934:14);

Stone, Knight and Starcke (1959); Belkin (1962:306-307); Nye and McGregor

(1964:11).

Mansonia iracundus of Miller (1950:44); Miller and Phillipps (1952:23-24).

Taeniorhynchus (Coquillettidia) iracundus of Edwards (1924:366).

Taeniorhynchus iracundus in part of Graham (1939:212).

FEMALE (Miller and Phillipps 1952:fig.11). Wing: 4.0 mm. Proboscis:

2.4mm. Forefemur: 1.8mm. Abdomen: about 2.7mm. As described by

Belkin (1962:306); superficially very similar to tenuipalpis from which, how-

ever, it can be readily distinguished by the following conspicuous features.

Size smaller; base of lower surface of vein Sc with broad scales only; apn and

ppn with distinct narrow scales; stp scales not extending above bristles to base

of pra; abdominal sternites dark-scaled in the middle. Claws with only 1 or 2

long basal spicules on outer surface. Female genitalia not studied.

MALE. Essentially as in the female except for sexual characters; differing

markedly in the latter from tenuipalpis as follows. Palpus not upturned, only

Slightly longer than proboscis; palpal segment 5 slightly shorter than segment

4; apex of segment 3 and all of 4 and 5 with numerous long hairs. Antenna much

shorter, less than 0.8 of proboscis; segments 12 and 13 together less than to-

tal length of basal flagellar segments, segment 13 distinctly shorter than seg-

ment 12.

Belkin: Culicidae of New Zealand 719

MALE GENITALIA (Belkin 1962:fig. 196; Nye and McGregor 1964:fig. 7).

As figured and described by Belkin (1962), with the following diagnostic features

and corrections. Segment IX: A small, well-defined tergite lobe bearing 2-5

bristles; sternite not as broad as in tenuipalpis but widely separating sidepieces,

sometimes with 1 bristle. Sidepiece simple, without any indication of lobe;

mesal membrane not developed but this area not as strongly sclerotized as in

tenuipalpis. Claspette with sternal process much shorter than in tenuipalpis,

not projecting beyond level of middle of aedeagus; apex with 1 long heavy,

apically truncate spiniform and 1, 2 apically attenuate thickened hairs. Clasper

with a conspicuous submedian outer lobe in an addition to a poorly defined dor-

sal lobe which extends from just beyond middle to apex; a conspicuous basal

dorsal ledge. Aedeagus short and widened subapically, poorly sclerotized on

dorsal surface, apex with 2,3 small teeth on each side dorsomesad; conspicu-

ous ventrolateral denticles extending from base to near middle. Paraproct with

about 5 large marginal spines and numerous smaller denticles on lower surface;

cercal setae usually 2, 3.

PUPA (fig.17). Abdomen: 3.2 mm. Trumpet: 0.76 mm (without decid-

uous apex). Paddle: 0.89 mm. Chaetotaxy as figured; diagnostic characters

as in the key; all normal culicine hairs present on cephalothorax and abdomen

except 2,3-VII (2-VII sometimes represented by alveolus), hair 1-IX present.

Cephalothorax lightly pigmented except for darkened dorsal surface; metano-

tum moderately pigmented. Trumpet very darkly pigmented on tracheoid;

bulb moderately pigmented; neck markedly narrowed, less than 0.5 of bulb

width, lightly pigmented and with smooth apical margin; deciduous apex lacking

in all Specimens. Abdomen uniformly moderately pigmented; hair 9-VIII

cephalad of caudolateral angle and distinctly mesad of lateral margin. Paddle

uniformly moderately pigmented, lighter than abdomen; both outer and inner

margins subequally produced on apex.

LARVA. Unknown.

SYSTEMATICS. Although superficially very similar to rohit danse: Ci ara-

cunda is a very distinct species whose closest relative appears to be linealis

(Skuse, 1889) from Australia. These 2 species form the distinct iracunda group

which is very different from the dominant crassipes group of the palaeotropics

and whose closest relative appears to be the holarctic perturbans group on the

basis of general adult morphology, pupal trumpet and pupal chaetotaxy.

BIONOMICS. Living pupae and pupal exuviae of iracunda have been found

in association with tenuipalpis in shallow margins of ponds and lakes choked

with vegetation. As with tenuipalpis, no larvae were recovered from these

areas. Graham's report (1939:212) of larvae from ''sunny sheltered creek-beds

with a temperature of 850F, and heavily charged with strong-smelling decaying

vegetation" at Spirits Bay is undoubtedly erroneous. Unfortunately, the larvae

misidentified as iracunda by Graham could not be located in any museum in New

Zealand and the identity of this species cannot be determined until additional

material from this locality is obtained.

C. iracunda appears to be much more abundant than tenuipalpis. Both

sexes rest in vegetation in the breeding sites and adjacent banks. Females

were collected biting in these areas as well as in gullies a considerable dis-

tance from the breeding sites. Apparently this species may occasionally enter

buildings. According to Graham (1939:212), iracunda is a persistent night biter

in scrub and bush areas in Auckland Province and is especially troublesome

around farms where it pesters the stock.

The presence of iracunda on Great Barrier and Mayor islands is of con-

siderable interest and indicates that this species has a considerable flight range

80 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

and is capable of surviving in very small populations.

DISTRIBUTION (fig. 35). Material examined: 342 specimens, 11 0%, 211 9,

120 pupal skins; 4 individual pupal rearings.

NEW ZEALAND. North Auckland: Spirits Bay, 19-24 Feb 1929, W.R.B.

Oliver, 52 [WELL]. Te Kao, 28 Feb 1919, D. Miller, 12 'USNM]; 1 Mar 1919,

D. Miller, 3¢[BMNH]. Kaitaia, Rangitahi bush, 6 Feb 1917, D. Miller, 2 2

[BMNH]; Kaitaia Hotel, on wall, 16 Feb 1965, F (NZ 204B), 19 [UCLA]. Hor-

eke, bush, 18 Jan 1917, D. Miller, 19 [BMNH]. Pahia, 1Jan1953, K.P. Lamb,

12 [AUCK]; 23 Jan 1946, G.L. Bissett (847), 12[NELS]. Kawakawa, bush

stream, 13 Jan 1919, D. Miller, 2c, 22[BMNH]. Ruakaka, 1 miS, 20 Dec

1963, BS (NZ 36), 1 p[UCLA]. Bethells Road Swamp, 28, 29 Dec 1963, BFS

(NZ 57A,57B), 1 po’ (57B-11), 3 p? (57B-10, 12, 13), 50%, 509, 115 p[UCLA];

10 Jan 1965, F (NZ 200A), 20°, 92[UCLA]. Quarry tributary of Waitakere

River, edge of Cascade Park, 28 Dec 1963, BF (NZ 59), 52 [UCLA]. Cascade

Park, Waitakere stream, 20 Feb 1964, F (NZ 177), 99 [UCLA]. Bethells Road,

Cascades turnoff, 15 Nov 1964, F (NZ 195A), 1c, 139[UCLA]. Waitakeres,

2,25, 26 Feb 1950, M. Knight, 3 2[AUCK], 29[UCLA]. Auckland Watershed

Reservoir, 26 Feb 1944, G. Chamberlain, 31 2 [WELL], 32 2[UCLA]. Auck-

land, 10 Jan 1924, G.H. Hopkins, 1 2 [BMNH].

Great Barrier Island: Warrens Creek, 16 Feb 1919, D. Miller, 39 [BMNH].

Mayor Island: 28 Nov 1942, S.A. Rumsey, 1 2? [AUCK].

Wellington: Ohakune, 15 Dec 1922-15 Jan 1923, T.R. Harris, 19 [BMNH].

Westland: Lake Brunner, 2,4 Feb1922, A.L. Tonnoir, 29 [NELS]. Hercules

Mt vicinity, Harihari, 7 Feb 1964, S(NZ), 12 [UCLA]. Lake Mapourika, 7

Feb 1964, S (NZ 112), 12 [UCLA]. Waiho, 16 Jan 1922, A.L. Tonnoir, 1 ?

[NELS], 1 2 [CANT].

New Zealand, locality not specified: Holotype ¢ (42-55) [BMNH]; 2 9°

[WELL]; 10 2 (683) [NELS].

TRIBE AEDINI

The 6 species of the tribe known to occur in New Zealand are readily rec-

ognized from other Culicinae by the following combination of characters. In

the adults postspiracular bristles are present. In the pupae, hair 9 on seg-

ment VIII is located at the caudolateral angle and paddle hair 1-P is present.

The siphon of the larvae is distinctive in the presence of a single pair of sub-

ventral tufts and the ventral brush is strongly developed. A general account of

the tribe is given by Belkin (1962:317-318).

SYSTEMATICS AND BIOGEOGRAPHY. The Aedini are better represented

in New Zealand than any other tribe of the Culicinae: by the monotypic endemic

genus Opifex, and 5 species in 4 subgenera (1 endemic) of the widespread genus

Aedes.

Opifex fuscus probably has retained more important primitive features

than any other living representative of the tribe and shows many unique derived

features in all stages (see below). It has the widest distribution in the New

Zealand area, occurring in the Kermadecs, the main islands of New Zealand

and the adjacent islets. It was probably the earliest member of the tribe to reach

New Zealand but there is no indication of geographical differentiation in this

species.

The subgenus Finlaya is represented in New Zealand by notoscriptus which

Belkin: Culicidae of New Zealand 81

has awide distribution in the Australasian area from the Moluccas to Australia,

New Caledonia and the Loyalty Islands. Since it is a semi-domestic species,

apparently recovered several times on ships from Sydney in New Zealand ports

(Graham 1939:212) and‘is still largely confined to the vicinity of these areas, I

consider that it was probably introduced to New Zealand during and following

colonization.

The remaining 4 species of Aedes are indigenous to New Zealand in my

opinion, and all except australis are endemic. A. australis is the only member

of the subgenus Halaedes and occurs elsewhere along the coasts of southeastern,

eastern and western Australia, Tasmania and Lord Howe and Norfolk islands.

Sharing many features with australis is the aberrant chathamicus, the only rep-

resentative of the endemic subgenus Nothoskusea, which is known only from

Chatham Island. A. chathamicus shows more primitive features than australis

and I consider that it was derived from a stock which also gave rise later to

australis. The immature stages of both chathamicus and australis show striking

Similarities with Opifex fuscus, especially in the terminal abdominal segments

and the head and abdominal chaetotaxy of the larvae, and the pupal trumpets,

paddles and general pattern of abdominal chaetotaxy. Although some of these

Similarities are undoubtedly due to convergence concomittant with the occupa-

tion of similar breeding sites, others are so close that it is evident that they

must be due to close genetic relationship. Of the 2 species, chathamicus shows

more similarities with Opifex and this suggests that it is probably an earlier

derivative than australis. It may have been derived at the same time as Opifex

fuscus; this is suggested also by the complementary distribution and the strik-

ing differentiation of the 2 forms in adult morphology, particularly in Opifex,

which unquestionably should be regarded as a genus distinct from Aedes. The

known distribution of australis in New Zealand is confined to the southern part

of South Island and Stewart Island. This species is sympatric with Opifex atleast

in eastern Otago. It is probable that australis is a more recent arrival in the

New Zealand area than chathamicus and Opifex; however, I disagree with

Mark's suggestion (in Nye 1962:33) that it is a recent introduction and consider

that this distribution is a natural one similar to the dispersals of australis to

Lord Howe and Norfolk islands.

The remaining 2 species of Aedes belong to 2 distinct sections of subgenus

Ochlerotatus (Marks and Nye 1963). A. antipodeus has been placed in the Vigi-

lax Section by Marks (1957; Marks and Nye 1963:54) on the basis of general

features of male genitalia and it appears to be most similar to inexpectatus

Bonne-Wepster, 1948 from the highlands of New Guinea on the basis of details

of male genitalia, scale pattern of adults and hair 2 of abdominal segments II-

VII of the pupa. However, in other pupal characters and larval features the 2

species do not show close affinity. This apparent relationship between a New

Zealand species and a species from the highlands of New Guinea is paralleled

by that of pervigilans and miraculosus in the genus Culex, according to Marks

and Nye (loc. cit.). A. subalbirostris appears to be allied to Australian species

of the Perkinsi Section and to 2 South African species of Ochlerotatus whose af-

finities seem to be also with this section (Marks and Nye 1963:59). According

to these authors this group is obviously an ancient one but insufficient know-

ledges of its components precludes even a suggestion as to whether albirostris

represents a derived or a primitive stock. All that can be said safely for the

present about the Ochlerotatus fauna of New Zealand is that it consists of 2 very

distinct species belonging to 2 ancient groups; that antipodeus has a wide dis-

tribution on North Island and South Island and occurs on some outlying islands

82 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

(Little Barrier and Great Barrier); that subalbirostris is apparently restricted

to South Island from Dunedin district to Invercargill and that it is sympatric

with antipodeus in this area; and that both species seem to have a wide range

in breeding sites and can tolerate at least low salinity.

It is interesting that the indigenous aedine fauna of New Zealand consists

entirely of ground pool and rock hole breeders. Since plant container habitats

(treeholes and leaf axils) are available and are utilized by other indigenous

forms (Maorigoeldia and Culex), this suggests that container breeding in Aedes

is a specialization which arose relatively late, after the dispersal of the genus

to New Zealand.

KEYS TO GENERA, SUBGENERA AND SPECIES

Adults

- Erect scales absent on head, instead numerous short curved bristles;

scape of antenna a distinct segment proximad of torus (Opifex).

De aes hs fuscus

Erect scales present on head: scape of antenna not recognizable as a

Str Seer LACORS. oe ke es es es a ak a de ewe 2

2(1). Decumbent scales of vertex of head largely broad; mesonotum with a

conspicuous pattern of narrow Silvery and golden-scaled lines in-

cluding a complete continuous lateral prescutal-fossal-posterior dor-

socentral line; scutellum with broad aid or light golden scales

(Pinlaya) i. i ice ce es ered MOLOSeriLptus

Decumbent scales of gure frecia’ narrow; mesonotum without continu-

ous lateral line as described above; scutellum with narrow scales on-

3(2). All tarsi with some distinct white-scaled basal bands; female cercus

long, narrow and projecting (Ochlerotatus) .. . 4

None of the tarsi with distinct white-scaled basal bands: female cercus

SHOTL, OrOad and NOL PrOjeCline 9 ee ve Oe wee og ee kent ie PO

4(3). Mesonotum with conspicuous narrow incomplete golden-scaled lines and

patches contrasting with dark-scaled background; vertex of head

with pair of ae submedian patches of narrow dark scales

ee 20. antipodeus

Mesonotal scaling more or less norm, without conspicuous golden

lines or patches; all narrow decumbent scales of vertex pale.

.21. subalbirostris

o(3). Anterior lower mep bristles absent; ssp without scales; psp usually

without scales, rarely with a few narrow ones (Nothoskusea) .

es chathamicus

Anterior lower mep bristles present: ssp with a ine of broad light

scales; psp with numerous broad scales (Halaedes). . . 19. australis

Male Genitalia

T; Cercal setae of proctiger of 2 types, short distally, long proximally;

aA)

a2).

4(2).

5(4).

4(3).

5(3).

Belkin: Culicidae of New Zealand 83

sidepiece without long bristles distally (Opifex) ... . . 17. fuscus

Cercal setae of proctiger all short; sidepiece with long bristles distally

(ROGGR) Gee ei ee a ey hia

Claspette short, with numerous bristles or foliform setae and without

apical specialized appendage or thickened bristle. ... 3

Claspette longer, with apical specialized appendage or thickened bristle

Basal tergomesal margin of sidepiece and claspette without foliform se-

tae (Nothoskusea). ... : ~ 2 418. Chathamicus

Basal tergomesal margin of sidepiece and claspette with foliform setae

CHeINCUGR) i. faye coe ee ee as oki a 2 es eS

Sidepiece without basal or apical tergomesal lobe, its sternomesal mar-

gin with line of lanceolate-tipped setae (Finlaya) . . .22. notoscriptus

Sidepiece with a prominent basal or apical tergomesal lobe, its sterno-

mesal margin without lanceolate-tipped setae (Ochlerotatus). . .. 5

Sidepiece with a large prominant basal tergomesal lobe and without dis-

tinct apical lobe .. . . . . 20. antipodeus

Sidepiece with a small, flat basal tergomesal lobe and a prominent

ADIGA 1Q0G f cutcy ew le ee ee oe we Nea | ee Be oSetrie

Pupae

Hair 9-VIlI ventral in position, simple, short, similar to 9-II-VI (Opi-

OR) 00's 1. faseus

Hair 9-VII dorsal it in n position, markedly d different from 9 <T VI see

. Hairs 10,11-C usually all single (Finlaya) . ... . . .22. notoscriptus

At least hairs 10, 12-C always praneied . 2.6 a ea 8

. Paddle width at least 0.75 of total width of segment VIII; trumpet very

broad, its length not more than 2.0 width . .

Paddle width less than 0.75 of total width of segment ‘VII: trumpet

longer, its length at less 3.0 width (Ochlerotatus) .... og

Abdominal hair 1-I (float hair) usually with not more than 20 simple

branches; paddle narrower than entire segment VIII, hair 1-P usually

branched ‘(Nothodkuten): . .18. chathamicus

Abdominal hair 1-I (float hai) es more ‘than 40 penipound branches;

paddle distinctly wider than eecrratns VIII, hair 1-P usually single

(Halaedes) 2 6); vu 4. 1 australis

Abdominal hair 2-III- VII sae ane submedian in position; hair 5-IV, V

usually single .. Bose ee, antipodeus

Abdominal hair 2-III- vl en in apical 0. 2 of seemente: hair 5-IV, V

usually double ... . : Fe Mae oni so subalbirostris

Larvae

Saddle occupying less than 0.5 of circumference of anal segment; gills

gua),

3(2).

4(1).

5(4).

1902.

Contrib. Amer. Ent. Inst., vol. 3, no. 1,.1968

tiny, rounded; abdominal hair 6-I-V usually at least triple, hair 7-I

usually at least double (Opifex, Nothoskusea, Halaedes). , 2

Saddle occupying more than 0.5 of circumference of anal Bdoment® anal

sills moderate, tapered or pointed distally; abdominal hair 6-I-V

usually double, hair 7-I usually single (Finlaya, Ochlerotatus). . . 4

Metathoracic pleural group (9-12-T) very small, longest hair less than

0.5 of long hairs in mesothoracic pleural group; pecten reduced to a

few teeth; hair 6-S very long, single (Opifex) . ... . . 17. fuscus

Metathoracic pleural group at most moderately reduced, longest hair

more than 0.5 of long hairs in mesothoracic pleural group; pecten

teeth numerous; hair 6-S moderate, branched (Aedes) ...... 3

Ventral brush with 9 pairs of hairs; pecten teeth in a continuous series

from near base of siphon; hair 9-T strongly developed, with many

branches, subequal to 9-M (Nothoskusea) . . . . . .18. chathamicus

Ventral brush with 7 pairs of hairs; pecten teeth largely absent from

base of siphon; hair 9-T weak, with few branches and much smaller

Pe Se ee) ke ea hs ee we ee trae

Head hair 1-C strongly thickened, sharply hooked mesad apically; ven-

tral brush with only 6 pairs of hairs, all on grid; siphon short, index

about 2.0; anal gills longer than saddle (Finlaya). . .22. notoscriptus

Head hair 1-C moderately thick but not sharply hooked mesad apically;

ventral brush with at least 7 pairs of hairs on grid and at least 1 de-

tached hair proximad of grid; siphon moderate, index more than 3.0;

anal gills shorter than saddle (Ochlerotatus). .......... 5

Siphon with pecten teeth evenly spaced; head hair 6-C single, 5-C usu-

ally at least triple .... » s 44 628, antipodeus

Siphon with 1 or more distal pecten teeth ‘widely spaced; head hairs 5,

oC ligually- both Gouna Go Ga Poe es CO eae, subalbirostris

Genus OPIFEX

Opifex Hutton, 1902:188. TYPE SPECIES: *O. fuscus Hutton, 1902; mono-

basic.

17. Opifex fuscus Hutton

Figs. 2,18, 37, 38

Opifex fuscus Hutton, 1902:188-189. TYPE: *Lectotype © (I- 290), Wel-

lington, N. Z., G. V. Hudson, marked as holotype; PRESENT SELEC-

TION, Hutton did not specify a type specimen in the original description

[CANT ].

Opifex fuscus of Edwards (1921:73-74; 1924:367; 1926a; 1932:127-128); Miller

(1922; 1950:45); Kirk (1923); Wood (1929); Taylor (1934:15); Graham (1939:

213); Lee (1944:47); Knight and Chamberlain (1948); Miller and Phillipps

(1952:11); Marks (1958); Stone, Knight and Starcke (1959); Dumbleton (1962);

Belkin (1962:322-324); McGregor (1963; 1964; 1965); Nye and McGregor

(1964:11-14); Haeger and Provost (1965).

Belkin: Culicidae of New Zealand 85

As described and figured by Belkin (1962:322-324) and as figured here;

with the following additions and diagnostic features.

FEMALE (fig.18; Miller 1922:figs.7-10; Miller and Phillipps 1952:fig.

16). Wing: 4.9mm. Proboscis: 2.25 mm. Forefemur: 2.0 mm. Abdomen:

about 3.0 mm. A medium-sized to large, stocky, gray-pruinose dark mosqui-

to with scanty scaling and uniformly slightly infuscated wing membrane. Head:

Interocular space about 0.5 as wide as high, with bristles to its lower margin;

frons high and broad above antennae, distance between inner articular points of

Scapes of antennae about 2.0 of their distance from lower margin of eye.

Vertex without erect scales, instead with numerous curved attenuate hairs.

Palpus as figured, about 0.28 of proboscis; apical segment (4) club-shaped and

with invaginated sense organ within apex of club. Antennae separated at base

by distinctly more than diameter of one torus; scape a distinct segment through-

out; torus moderately wide and short, its width about 1.6 of its length anda

little less than 2.0 of maximum width of flagellar segment 1; flagellar segment

1a little more than 1.5 length of segment 2; segment 3 alittle longer than 2 and

Slightly shorter than 4, remainder of flagellar segments subequal; flagellar

whorls extremely poorly developed, usually with only 6 hairs, none of them

longer than the segment except on 2-4. Proboscis (particularly lower surface),

palpal segment 4 and flagellar segments 4-13 with dense vestiture of short fine

hairs. Thorax: Mesonotal bristles very numerous, essentially as in chathami-

cus but shorter, heavier and without the concentration and elongation in supra-

alar and prescutellar areas; parascutellum without bristles. Scutellar bristles

not numerous, long ones confined to apical margin of lobes. Pleural bristles

as figured, numerous but relatively short except for some thickened ones; mid-

dle and lower mep bristles numerous. Legs: Foreleg markedly shortened;

bristles on all segments thickened, spiniform but short; plantar surfaces of all

tarsi with moderate vestiture of thin hairs projecting beyond scales. Claws of

all legs large, those of foreleg a little longer than others; members of a pair

equal; all with an inner subbasal long, slender straight tooth; spicules not de-

veloped on outer surface of base. Wing: As figured; vein Sc ending on C well

before furcation of Rs, its base ventrally with a large patch of bristles. Dor-

sal surface of veins Sc, Rs, R2+3, Rg, R3 and M with plume scales, those on

Sc broad; remaining veins with small squame scales. Alula with long margin-

al row of very long, narrow, striated hairlike scales except at base; upper cal-

ypter with complete marginal row of long bristles. Abdomen: Scaling very

sparse; tergite I with scales; sternite I partially sclerotized, without scales

or hairs. Hairs numerous and widespread on all tergites and sternites II-VII

but short except laterally.

FEMALE GENITALIA (fig.18). As figured. Completely retracted into

segment VIII, even cerci invisible at rest. Tergite VIII about 0.65 length of

sternite, with a few scalesand many bristles, fully exposed; sternite VIII strong-

ly compressed, without scales, with numerous bristles and with dense hydrofuge

short hairs distally, and with a conspicuous densely spiculose invaginated cau-

dal membrane. Tergite [IX represented bya pair of dorsolateral lobes separated

middorsally by membrane, apex of lobes with setae chiefly on mesal surface.

Insula and sigma continuous, narrow; latter joined to cowl; insula with 2-4 se-

tae on each side of midline. Cowl very broad laterally, with slight lateral atrial

expansion; broad ventrally where it is broadly joined to base of postgenital

plate. Tergite X not developed. Postgenital plate poorly sclerotized, triangu-

lar and with broad base; its ventral surface with numerous setae from base,

setae longer on apex. Cercus moderate, broad and with rounded apex; setae

86 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

numerous on outer surface.

MALE (figs. 2,18; Dumbleton 1962:fig.2; Miller 1922:figs.1-3,5; Miller

and Phillipps 1952:fig.17). Essentially as in the female but with remarkable

sexual differences as follows. Palpus about 0.8 of proboscis, pendent. Anten-

nal torus and flagellar whorls not enlarged; flagellar segments 2-4 with a dor-

sal basal swelling from which arises a greatly enlarged spiniform developed

from one of the whorl bristles, shortest on segment 3 and longest on 4. Fore-

tibia shortened and swollen, with a conspicuous fringe of long narrow scales

on ventral surface and dense vestiture of long apically-curved hairs near apex;

all tarsal segments with scaling confined to dorsal surface, plantar surfaces

with very conspicuous hydrofuge fringe composed of densely packed hairs of

varying size, the largest with curved apex; all tarsal segments thickened, seg-

ments 2-5 shortened, segment 3 with a long curved apical bristle. Midleg

elongated, particularly tarsal segments 1 and 2; hydrofuge fringe as on fore-

tarsus; tarsal segments 2-4 with long curved apical bristle. Hindleg with tar-

sal segment 1 a little longer than tibia; hydrofuge fringe as on foretarsus and

apical bristles as on midtarsus. All leg segments with scaling restricted and

with numerous fine hairs. All claws without tooth; those of foreleg tremen-

dously enlarged, longer than segments 3-5 combined. Abdominal scaling more

restricted than in female.

MALE GENITALIA (Belkin 1962:fig.207; Nye and McGregor 1964:fig. 8;

Dumbleton 1962:fig.1; Miller 1922:fig.6). Segment VIII markedly shortened,

only slightly retracted. Segment IX well developed and strongly sclerotized,

long but narrow; tergite without lobes but with submedian patch of numerous

short hairs; sternite with distinct basal and distal sclerites, without hairs.

Sidepiece with long hairs present only on dorsal surface in basal half near the

conspicuous lateral angle, remainder of surface, particularly side, with dense

hydrofuge pile of short curved hairs; mesal surface with extensive membrane

from base to apex. Clasper heavily sclerotized; short; strongly swollen and

with small rounded process at base, slender beyond; spiniform very heavy,

deeply inserted. Claspette a long simple sclerotized plaque in mesal mem-

brane; with short simple hairs distally. Aedeagus long, simple. Paraproct

with a single simple or serrated heavy spine. Cercal sclerite strongly devel-

oped, setae numerous, short distally, long proximally.

PUPA (Belkin 1962:fig.207; Knight and Chamberlain 1948:fig.24; Miller

1922:figs. 20-24). Abdomen: 4.04 mm. Trumpet: 0.62 mm. Paddle: 1.3mm.

In general similar to chathamicus and australis but differing markedly in the

following important features. Cephalothoracic middorsal ridge very strongly

developed. Trumpet short, extremely broad, index about 1.1; strongly com-

pressed, anterior margin straight, posterior curved; pinna terminal, not

markedly oblique. Abdominal hair 1 a simple hair with a few branches, nota

distinct float hair; hair 3-III caudad of level of hair 5; hair 5-IV, V usually sin-

gle; hair 9-VII small, single, similar to 9 on preceding segments, inserted on

ventral surface near caudolateral margin. Paddle distinctly wider than entire

width of tergite VIII, paddle hair 1-P usually single.

LARVA (Belkin 1962:fig.208; Nye and McGregor 1964:fig.9; Miller and

Phillipps 1952:fig.18; Lee 1948:fig.28; Miller 1922:figs.11-19). Head: 0.98

mm. Siphon: 1.07 mm. Anal Saddle: 0.43 mm. In general similar to chat-

hamicus and australis; with the following diagnostic features. Head hairs

4,6-C and 12,13-C far forward, 7-C very small. Antennal hair 1-A minute,

slender, spiniform. Thorax with mesothoracic and metathoracic hairs not

markedly approximated. Some of the prothoracic hairs moderately developed

Belkin: Culicidae of New Zealand 87

and other branched. Metathoracic pleural group (9-12-T) very small, with all

hairs single, at about level of hair 8-M but in normal position, far removed

distad. Abdominal hairs mostly weak; hair 6-VI relatively strongly developed,

not markedly different from 6-V. Comb in a triangular patch of 70 or more

scales; scales very small proximally, progressively larger distally, apical scales

frequently long thornlike spines. Siphon index usually about 2.0; Somewhat

swollen in middle, not strongly narrowed apically; acus not developed; trachea

greatly expanded in siphon, very dark; pecten represented by a few slender,

simple, widely-spaced spines; hair 1-S large, beyond middle of siphon and

distinctly ventrad of pecten; hair 6-S very long, barbed. Anal segment without

any indication of acus; saddle moderately well developed, covering most of dor-

sal surface of anal segment; hair 1-X on saddle; ventral brush usually with 8

pairs or 17 hairs. Anal gills extremely short, rounded, the 2 pairs usually

subequal.

SYSTEMATICS. Opifex fuscus is undoubtedly derived from the same stock

as A. chathamicus and A. australis. The first 2 species have retained in var-

ying degree and number many features which I consider to be primitive for the

subfamily Culicinae and, of course, possess many derived characters as well.

The more striking features in both instances are found in fuscus and for this

reason it is retained in a Separate monotypic genus.

The most important primitive features, unique in the tribe Aedini, exhibi-

ted by fuscus are: (1) in the adults a fully developed scape of the antenna, and

setae instead of erect scales on the vertex of the head; (2) in the pupa, hair 9-

VII similar to 9-II-VI. Both sexes have strongly developed bristles on the base

of vein Sc; this apparently primitive feature is known elsewhere in the tribe

Aedini only in the female of chathamicus (reduced or absent in the male of this

species and absent in both sexes of australis). Its larva appears to have the

most primitive pattern of the phylad in the following: (1) metathoracic pleural

group removed from the mesothoracic, (2) hair 6-VI of the same order of mag-

nitude as 6-V, and (3) anal saddle relatively well developed. The female has

the least differentiation of the flagellar segments and the shortest bristles in

the flagellar whorls.

The most conspicuous derived features in fuscus are: (1) in both sexes,

the development of a dense hydrofuge vestiture on the tarsi, the shortening of

the foreleg, and an invaginated pit on the apex of the palpus, (2) in the male,

specialization of flagellar segments 2-4 and legs, including tremendous en-

largement of the foreclaws, and hydrofuge vestiture of the genitalia, (3) in the

pupa, the development of a strong middorsal cephalothoracic ridge and the re-

duction of the float hair on abdominal segment 1, and (4) in the larva, reduction

of metathoracic pleural group and pecten. The derived features of the adults

and of the pupa appear to be all related to the unusual method of mating and

walking on the water surface in this species.

There is considerable variation in all stages of this species and particu-

larly in the larva (size, comb and pecten) but there is no indication of allopatric

differentiation. Even the isolated remote population on Kermadec Islands falls

within the range of variation in the populations on the main islands of New Zea-

land. Probably much of the variation noted is due to differences in food supply

in the breeding sites, as indicated by marked differences in size of the imma-

ture stages and adults from different sites.

BIONOMICS. Opifex fuscus has been found breeding primarily in brackish,

saline and hypersaline rock pools above high tide level. I collected it (NZ 65)

in a hole in a dead pohutukawa trunk projecting horizontally out over the shore

88 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

above high tide level. Larvae have also been taken in a fresh water stream

(Dumbleton 1962:24) and in a horse trough (L.C. Pilgrim in Nye and McGregor

(1964:12). |

The remarkable biology of Opifex fuscus and particularly its mating behav-

ior have received considerable attention (Miller 1922; Kirk 1923; Edwards

1926; Marks 1958; McGregor 1964,1965; Haeger and Provost 1965). Eggs are

laid primarily in small cracks and crannies above the water surface but in the

damp zone around the margins of rock pools. More than half of experimentally

dried eggs hatched after 6 months. The aquatic cycle is apparently quite long

in nature since at room temperature (mean 16.2°C) and normal day lengtha

mean period of nearly 24 days was required from the beginning of the second

larval instar to the emergence of adults, with more than 14 days for the fourth

larval instar. Experiments similar to those of Woodhill (1936) with Aedes aus-

tralis indicate that Opifex fuscus has a great salinity tolerance but not as ex-

treme as the former species (McGregor 1965).

Haeger and Provost (1965) established a colony of Opifex fuscus and main-

tained it at a rearing temperature of 259-279C. In the laboratory more pre-

cise observations on the biology of this species were made but theyare not al-

ways in agreement with field observations. The females are autogenous and

start to lay eggs 6 days after emergence. Both males and females can emerge

from the pupal skin unassisted but this seldom if ever happens in nature. Males

spend most of their time on the water surface hunting and competing for sur-

facing pupae. A male captures a pupa with its forelegs and foreclaws and at-

taches its claspers and part of the sidepiece to the middorsal cephalothoracic

ridge of the pupa. According to McGregor (1965:9), in nature when an adult

male captures a male pupa it releases it within 5 seconds of capture. In the

laboratory, however, Haeger and Provost (1965:26) found that the attending

male does not recognize the sex of the emerging adult until genital contact is

established. When the adult begins to emerge the attending male quickly in-

serts its genital segments into the thoracic region of the pupal skin and ap-

presses his genitalia to the side of the abdomen of the emerging adult. When

contact is established with the cerci of an emerging female, genital connection

is effected rapidly and the male runs for objects on the side of the container,

dragging the dangling female.

Populations of Opifex fuscus may reach very high densities, the surfaces

of some rock pools being almost covered by swarms of adults, mostly males.

On South Island apparently only larvae are found during the winter months but

on North Island and the Kermadecs adults probably are present throughout the

year.

Females are said to attack man at night (Graham 1939:213) and to produce

a most painful bite (Miller and Phillipps 1952:11).

DISTRIBUTION (figs.37, 38). Material examined: 1550 specimens; 335,

169 2, 783 larvae, 263 pupae; 69 individual rearings (51 larval, 11 pupal, 7 in-

complete).

KERMADEC ISLANDS. Raoul Island: Station, 21-30 Sept 1962, G.A.

Samuelson, light trap, 39 [BISH], 19 [UCLA]. Lava Pt., 7 Oct 1962, G.A.

Samuelson (130), littoral saline and brackish pools, 17, 27 L[BISH], 10,

10 L[UCLA]. Locality not specified, 9 Oct 1908 (12), 4 o% [CANT].

S. Meyer Island: 15 Oct 1962, G.A. Samuelson (155), shallow littoral

brackish pool, 102 ¢, 19, 4L, 1 P[BISH], 20°, 4L, 1 P[UCLA].

Kermadec Islands, locality not specified: Aug 1908, 1 o [WELL].

NEW ZEALAND. Three Kings: Great Island, E.G. Turbott, 17 Apr 1946,

Belkin: Culicidae of New Zealand | 89

3 ¢, 6 May 1946, 29, 11 Jan 1951, 1 2; Tasman Valley, 5 Jan 1953, G. Archey,

1 9 [AUCK; Marks 1958:20].

North Auckland: Kaitaia, 10-11 Oct 1924, 39 [WELL]. Russell, 30 Mar

1925, 1? [WELL]; Russell Beach, 1928, D. Miller, 1 co [LOND]. Pahia, 2

Mar 1949, K.P. Lamb, 19[AUCK]. Ocean Beach, Whangarei Heads, 31 May

1964, F (NZ 187), 21po% (187B-105, 109), 71p? (187B-102-104, 106-108, 111),

1 po (187B-110), 2 1p (187B-101, 112), 123 o (187A), 90 2 (187A), 11 L (187B),

16 P (187B) [UCLA]. HenIs. [NELS; Dumbleton 1962:24]. Whangaporoa Pen-

insula, 15 Mar 1949, D. Spiller, 1 9 [AUCK]. Milford, 4 Mar 1927, D.H. Gra-

ham (23-19), 12 [LOND]. Rakino Island, 24, 25 Jan 1964, F (NZ 172, 173), 1,

13 L, 1 P [UCLA]. Waiheke Island, 1 Mar 1927, D.H. Graham (23-18), 1 9

[LOND]. Eastern Beach, 8 Jan 1945, G.L. Bissett (520), 1 L [NELS]. Little

Huia, 30 Dec 1963, BF (NZ 65), 9 lpo (65-102, 103, 105, 107-109, 112, 114,

115), 71p (65-104, 106, 110, 111, 113, 117, 119), 1 po (65-101), 11p (65-118),

60, 179) 652, 31 PPUCUAT.

Mokohinau Islands: Burgess Island, 24 Aug 1955, R.A. Harrison, 10, 39

[AUCK]. Lizard Island, 23 Aug 1955, R.A. Harrison, 7 o [AUCK]. On board

ship, 11 Mar 1964, S (NZ 135), 8 9 [UCLA].

Great Barrier Island: Mouth of Te Wairere stream, 26 Oct 1963, S (NZ

17), 51po (17-101, 104, 105, 109, 111), 3 1p (17-102, 103, 106), 1 1p (17-107),

7, 69, 60 L, 24 P[UCLA].

Cuvier Island: 24 Oct 1963, S (NZ 14), 6 lpo" (14-102, 103, 106, 107, 110,

111), 31p? (14-104, 105, 108), 1 p2 (14-101), 11p (14-109), 4c, 14 L, 8 P

[UCLA]; 13 Mar 1964, S (NZ 137), 1 lpo (137-103), 8 1p (137-104-106, 108,

109, 111,114,119), 6 po (137-110, 112, 113, 115-117), 2 p? (137-101, 107), 21p

(137-102, 118), 521 L, 112 P [UCLA].

South Auckland: Tauranga, 5 Apr 1945, C.W. Dixon (680), 4 o, 1 9 [NELS].

Wellington: Kapiti Island, 23 Dec 1943 (186), 3 L [NELS]. Somes Island,

19 Oct 1944, R.K. Dell, 40, 19, 4L, 3 P [WELL]. Wellington, G.V. Hud-

son, lectotype “ (I-290), paratype ? (I- 291) [CANT], 2 o [BMNH]; 1921, G.V.

Hudson, 2°, 59, 23 L, 1 P[BMNH], 1c, 3 L, 3 P [USNM]; (2364), 10, 19

[LOND]. Wellington, Breaker Bay, 16-25 Jan 1939, J.T. Salmon, 2 o%, 2 9

[WELL]; 23 Mar 1944, W.J. Phillipps (172), 1 L[NELS]. Wellington, Hough-

ton Bay, 6 Dec 1943, V. Brazendale, 2c, 22 [WELL]; 10 Dec 1943, Russell,

2% [WELL]; 30 Mar 1944, W.J. Phillipps (183), 1 L, 1 P [NELS]. Welling-

ton, Island Bay, 15 Jan 1925, 1c, 1? [WELL]; 18 Jan 1943, W.J. Phillipps,

29°[WELL]; 22 Nov 1943, V. Brazendale, 19 [WELL]; 21 Sept 1944 (414), 13

L, 4P [NELS]; 12 Dec 1953, J.C. Yaldwyn, 98 L, 17 P [UCLA]; 3 Jan 1954,

J.C. Yaldwyn, 230%, 11 9[UCLA]; 24 Jan 1957, E.N. Marks [Marks 1958:20];

Apr 1961 [Haeger and Provost 1965]. Wellington, Owhiro Bay, 9 Mar 1944,

W.J. Phillipps (125), 1 L [NELS].

Nelson: Cable Bay [Dumbleton 1962:24].

Marlborough: Kaikoura, 11 Feb 1922, Archey, 1 o [CANT], 1961 [McGre-

gor 1964;1965]. Goose Bay, 4 Feb 1925, A.L. Tonnoir, 7, 3 9 [CANT].

Canterbury: Banks Peninsula, near Akaroa lighthouse, R.L.C. Pilgrim,

in horse trough [Nye and McGregor 1964:12]; Murray's Mistake, in fresh-water

stream [Dumbleton 1962:24]; spray zone, 1961 [McGregor 1964; 1965].

Otago: Martin's Bay, 1-4 Feb 1955, R.R. Forster, 9%, 19 [CANT]. Ota-

go Peninsula, Cape Saunders, 24 Feb 1918, 12 [AUCK; Marks 1958:20], all

stages [Nye and McGregor 1964:12]; Sandfly Bay, all stages [Nye and McGregor

1964:12].

New Zealand, locality not specified: 1 Apr 1928, 1 L, 1 P [LOND]; E.S.

90 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

Gourlay (138), 1 L[NELS]; 3 Jan 1943, 1 9 [UCLA].

Genus AEDES

Subgenus Nothoskusea

1962. Nothoskusea Dumbleton, 1962:20. TYPE SPECIES: *Aedes (Nothoskusea)

chathamicus Dumbleton, 1962; original designation, monobasic.

18. Aedes (Nothoskusea) chathamicus Dumbleton

| Figs. 19-21, 37, 38

1962. Aedes (Nothoskusea) chathamicus Dumbleton, 1962:20-24. TYPE: *Holo-

type o, bred from pupa, south of Point Weeding, Chatham Is., 18 Nov

1959, L.J. Dumbleton [NELS]. 7 |

Aedes (Nothoskusea) chathamicus of Stone (1963:175).

In general as described by Dumbleton, with the following additions and

corrections. The largest mosquito in the New Zealand area in the adult and

immature stages.

FEMALE (fig. 19; Dumbleton 1962:figs. 3,4). Wing: 6.0 mm. Proboscis:

2.8mm. Forefemur: 3.4mm. Abdomen: about 4.0 mm. Large dark species;

external morphology as figured. Head: Interocular space about as wide as in

Opifex and with short bristles and proclinate narrow golden scales nearly to its

lower border; frons high and broad above antennae, distance between inner ar-

ticular points of scapes of antennae greater than their distance from lower mar-

gin of eye. Vertex with sparse vestiture of decumbent or proclinate narrow

scales, pale golden in a longitudinal median line and in back, dark bronzy in

submedian anterior patch of variable extent; some broad decumbent pale gold-

en or yellowish scales laterally, extending back of orbital margin a variable

distance toward midline; erect scales numerous, all dark, very long, slender

bristlelike but with truncate irregularly or shallowly forked apex; sides with

scattered decumbent pale scales; upper orbital bristles in 2 or more irregular

rows, with a pair of longer differentiated ones at upper end of interocular

space; lateral orbital bristles in a regular single row; numerous shorter

bristles on sides and ventrally following erect scales; clypeus bulging, bare,

pollinose. Proboscis distinctly shorter than forefemur; labium entirely dark-

scaled; labella long, slender, without scales. Palpus about 0.23 of proboscis;

segments 1,2 and 3 indistinctly separated; 1 short and bare; 2 subequal to 1,

with short and long bristles and a fewsmall scales; segment 3 about 2.3 of seg-

ment 2, with short and long bristles and numerous scales; segment 4 subequal

to 1-3 combined, with short bristles and numerous scales; segment 5 a little

shorter than segment 1, spherical and bare; all palpal scales dark. Antennae

Separated at base by distinctly more than diameter of one torus; subequal to

proboscis in length; scape a narrow ring except for conspicuous extension at

mesal articular point with torus; torus moderate, width subequal to length and

only about 2.4-3.0 maximum width of flagellar segment 1, mesal surface with

numerous bristles and a few minute broad scales; flagellar segment 1 a little

more than 1.5 length of segment 2, with a few minute scales on mesal surface,

Belkin: Culicidae of New Zealand 91

hairs numerous, long ones in distal half confined to dorsal and lateral surface;

segments 2-13 progressively longer, segment 13 a little less than 2.4 length of

segment 2; flagellar whorls poorly developed, usually with only 6 differentiated

hairs, the dorsal, ventral and 2 outer long, the 2 inner very short, all pro-

gressively shorter on distal segments; segment 13 with apical process and with

about 3 moderately long subapical bristles (possibly rudiment of flagellar seg-

14, also noted in many other New Zealand Culicinae). Thorax: Mesonotal in-

tegument dark brown; scaling entirely narrow, very dense along lines of bris-

tles, Sparser in between, predominantly dark bronzy but with a more or less

distinct and variable pattern of light golden scales on anterior promontory,

usually scattered on acrostichal line with a patch near its middle and its pre-

scutellar end, on middle of lateral prescutal line, at mesal end of fossal line,

laterad of prescutellar bristles and above wing root, sometimes forming an in-

complete linear pattern. Mesonotal bristles very numerous, short except pos-

terior prescutellar and caudal supraalars; acrostichals in 2 widely separated

irregular lines; each dorsocentral line in 2 irregular rows; fossals numerous

and merging with anterior dorsocentrals and humerals; a more or less distinct

posterior fossal row; supraalars extremely numerous; prescutellars numerous;

parascutellum with a strong bristle. Scutellum with strongly differentiated

lobes; scales narrow as on mesonotum, midlobe with light golden scales in the

middle and dark bronzy laterally, lateral lobes with light golden scales only;

bristles very numerous, occupying most of surface of all lobes, varied in size,

extremely long caudad. Paratergite long, rather narrow, bare. Pleural integ-

ument brown and with darkened areas; pleural scaling very sparse, scales pre-

dominantly moderately narrow, semierect and light golden except on ppn where

they are dark, present on apn, ppn, ppl, psp, upper and lower posterior stp, be-

low bristles on pra, and middle of mep to upper mep bristles; bristles very nu-

merous, as figured, lower mep bristles absent. Legs: Coxae with large patch

of scales on anterior surface, scales all dark on anterior leg, dark on others

except for variable number of light scales at base; scaling of remainder dark

except for light extreme apices of femora and tibiae and base of hindtarsal seg-

ment 1. Femora subequal in length but midfemur a little longer than others;

forefemur with large patch of specialized light bristles on apex of anterior sur-

face. Tibiae subequal in length to femora of same leg; that of hindleg a little

longer than its femur; fore and particularly hindtibiae with strongly developed

apical combs; first tarsal segments progressively longer on fore, mid and hind-

leg and always shorter than corresponding tibiae; all tarsal segments with rows

of minute straight thornlike setae shorter than the scales on plantar surface,

particularly strong on first segments. Claws moderately large, subequal on

fore and midleg but distinctly shorter on hindleg, members of a pair equal; all

with short spicules on outer surface of base and with a rather heavy subbasal

tooth as figured. Wing: As figured; vein Sc ending on C well beyond furcation of

Rs, its base ventrally with a large patch of moderately long bristles. Dorsal

surface of veins Sc, Rs, R913, Ra, R3, M and distal part of 1A with plume

scales, narrow on all except Sc, remaining veins with squame scales. Alula

with a complete marginal row of long narrow scales except in basal 0.3; upper

calypter with complete marginal row of longer bristles except at base; lower

calypter bare. Abdomen: Predominantly dark-scaled; tergites II-VII usually

with patch of pale scales basolaterally, frequently more or less complete trans-

verse basal pale bands connecting lateral patches, more widely on anterior

segments; segment I without scales, its sternite not sclerotized but with 1-3

short bristles on each side, its tergite with very dense vestiture of very long

92 Contrib: Amer. Ent. Inst.,:vol..3, no. 1, 1968

and shorter hairs. Segments II-VII with numerous hairs on tergites and ster-

nites not confined to lateral and apical areas.

FEMALE GENITALIA (fig.19; Dumbleton 1962:fig.4). As figured here.

Almost completely retracted into segment VII. Segment VIII with only a few

scales usually developed; tergite about 0.5 of length of tergite VII; sternite not

markedly larger, about 1.25 of tergite VIII and 0.625 of sternite VI. Tergite

IX strongly developed, about 0.5 of tergite VIII, separated middorsally into 2

lateral sclerites, each with 5-12 setae distally. Insula and sigma continuous,

extremely narrow; latter joined to cowl; insula with 2-4 setae on each side of

midline. Cowl very broad laterally, with large atrial expansion, narrowed

mesad where it is joined to the broad apically rounded more lightly sclerotized

postgenital plate which bears numerous Setae distally; a small heavily sclero-

tized area at center of base of postgenital plate. Tergite X not developed. Cer-

cus broad, subequal in length to tergite IX, with numerous setae laterally and

apically.

MALE (fig.19; Dumbleton 1962:figs.5,6). Very similar to female, dif-

fering primarily in the following. Palpus about the same length as in the fe-

male, similar in structure except that segment 3 is a little longer and 4 a little

shorter and segment 2 has numerous scales. Antenna not markedly different

from female; torus not distinctly larger, slightly wider in proportion to width

of flagellar segment 1; flagellar segments 2,3 and 4 shorter and apical seg-

ments longer as figured. Pleural scaling sparser. Claws essentially as in the

female but slightly shorter and not as heavy, subbasal tooth more slender. Wing

shorter; basal ventral bristles of Sc not developed or very short. Abdomen

distinctly longer than in female and much longer than the wing; scaling of ab-

domen very sparse, light scaling indistinct; bristles heavier and more numer-

ous.

MALE GENITALIA (fig.20; Dumbleton 1962:fig.7). As figured here.

Segment IX very narrow; tergite with poorly developed lobes bearing about 10

thin bristles; sternite with a distal patch of more than 20 longer, heavier bris-

tles. Sidepiece with long hairs laterally and ventrally to apex; mesal mem-

brane poorly developed, narrow; tergomesal area with numerous short, heavy

simple bristles; sternomesal area with numerous short, thin simple bristles.

Clasper heavily sclerotized, sinuous in outline; spiniform short and heavy.

Claspette a simple prominent lobe with numerous simple short bristles primar-

ily on outer surface. Aedeagus very small, simple. Paraproct with a single

heavy apical tooth; cercal setae all similar, very short, usually 4-8.

PUPA (fig.20; Dumbleton 1962:figs.8,9). Abdomen: 5.5mm. Trumpet:

0.64 mm. Paddle: 1.28 mm. In general very similar to australis; chaetotaxy

as figured here. Cephalothoracic middorsal ridge normal. Trumpet short and

broad, index usually about 1.7; broadly flared, not distinctly compressed, both

anterior and posterior margins straight, pinna only slightly oblique. Abdomi-

nal hair 1-I usually with not more than 20 simple branches; hair 3-III caudad

of level of hair 5; hair 5-IV, V usually triple (2-4); hair 9-VII strong, distinct-

ly differentiated from 9 on preceding segments, at lateral margin of tergite.

Paddle distinctly narrower than entire width of tergite VIII, paddle hair 1-P

usually branched.

LARVA (fig.21; Dumbleton 1962:figs.10-13). Head: 1.15 mm. Siphon:

1.34 mm. Anal Saddle: 0.51 mm. In general very similar to australis; body

much longer but sclerotized parts not markedly larger; chaetotaxy as figured.

Head uniformly moderately pigmented except for darkened caudal areas, mar-

gin of foramen magnum and collar; head hairs 5,6-C both moderately strong,

Belkin: Culicidae of New Zealand 93

5-C usually double (1-3), 6-C single; 13-C usually double or triple, moderately

strong. Antenna concolorous with head capsule in basal part, slightly dark-

ened distad of hair 1-A; shaft slender but somewhat irregular, with incon-

spicuous fine spicules; hair 1-A usually with about 10 branches (8-15). Thorax

with mesothoracic and metathoracic hairs close together and far removed from

prothoracic. Prothoracic hairs all weak but several larger hairs with 3 or

more branches. Metathoracic pleural group (9-12-T) moderately large, dor-

sad of hair 8-M and separated from hairs 6,7-M by more than twice length of

their common basal tubercle, hair 9-T moderate, usually with 10 branches (7-

13). Abdominal hairs weak; hair 6-VI about 2.0 of hair 4. Comb in a large

patch of more than 100 minute broad scales fringed apically. Siphon character-

istically shaped as in the figure; index usually a little more than 2.0; trachea

greatly expanded in siphon, black; a distinct narrow attached acus present;

pecten in a continuous series or interrupted near the middle, with 12-30 simi-

lar teeth often extending to near middle of siphon length; hair 1-S beyond mid-

dle. Anal segment with a very small detached ventrolateral acus; saddle

largely dorsal but longer than in australis; hair 1-X close to saddle; ventral

brush usually with 9 pairs of hairs. Anal gills very short, rounded, dorsal

pair almost twice as large as ventral.

SYSTEMATICS. Aedes chathamicus, as indicated under O. fuscus, un-

doubtedly belongs to the same stock as the latter and A. australis. It possesses

a number of primitive features and lacks any very striking derivative features

of the type present in Opifex. It is tempting, therefore, to consider it as the

most primitive member of this phylad. However, it lacks the very striking

unique primitive features of Opifex (see) and therefore it does not seem prob-

able that the latter was derived from it. On the other hand, it is clear that

australis is a derivative from a chathamicus-like stock and in many important

features chathamicus suggests an intermediate step between fuscus and austra-

lis. I therefore submit that Opifex was the first segregate of this phyletic line,

followed by chathamicus and finally australis.

The most important primitive features of chathamicus appear to be: (1)

interocular space and frons wide, essentially as in Opifex, (2) antenna essen-

tially similar in the 2 sexes with relatively small torus and poorly developed

bristles in the flagellar whorls, (3) palpus essentially similar in the 2 sexes,

segment 5 present, (4) legs without special modifications, claws toothed and

with spicules, essentially similar in the 2 sexes, (5) numerous bristles on ab-

dominal segments, (6) female and male genitalia least modified in the phyletic

line, and (7) ventral brush of larva most strongly developed. All of these char-

acters except (1),(5), and (7) have been strongly modified in Opifex but these

are all associated with the unusual sexual dimorphism and mating habits of the

latter species.

The derived features of chathamicus, intermediate between Opifex and aus-

tralis, are: (1) in the adult, differentiation of flagellar segments and bristles in

flagellar whorls, (2) in the pupa, differentiation of hair 9-VI, and (3) in the lar-

va, association of metathoracic pleural group with hairs of mesothorax, reduc-

tion of hair 6-VI, and reduction of anal saddle. The other derivative features

of chathamicus are primarily in the reduction of the pleural scaling and the

elongation of the male abdomen.

BIONOMICS. The immature stages of chathamicus are known only from

saline or brackish rock pools at or just above high tide mark. No one other

Species of mosquito has been found associated with chathamicus. Adults of this

species have not been encountered in nature, all known specimens were reared.

94 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

DISTRIBUTION (figs. 37, 38). Material examined: 364 specimens; 25 0, 329,

211 larvae, 96 pupae; 61 individual rearings (12 larval, 37 pupal, 12 incom-

plete).

CHATHAM ISLANDS. Chatham Island: Rocky coast adjoining cliffs south

of Pt. Weeding, near Waitangi, 18 Nov 1959, L.J. Dumbleton, © holotype

[NELS]. Te Awarakau, near Waitangi, 25 Feb 1964, S (NZ 150,155), 41lpo

(155-106, 108, 122,140), 8 1p? (150-101, 105, 106,108; 155-107, 126, 127, 1477),

14po (150-102, 111; 155-104; 105, 109-112, 114, 118,123, 182,136,150), 23 p?

(150-107, 109; 155-101-103, 113, 115-117, 124, 125, 128-131, 135, 187, 138, 142-

146), 12 lp (150-103, 104, 110, 112,113; 155-119, 133, 134, 139, 141, 148, 149)

[UCLA]. Point Durham, rocky coast [Dumbleton 1962:21].

Subgenus Halaedes

1914. Caenocephalus Taylor 1914:700. TYPE SPECIES: C. concolor Taylor,

1914, New South Wales; monobasic. Preoccupied by Caenocephalus v.

der Wulp, 1898.

1962. Halaedes Belkin, 1962:328-329. TYPE SPECIES: Culex australis Erich-

son, 1842, Tasmania; original designation.

19. Aedes (Halaedes) australis (Erichson)

Figs. 37, 38

1842. Culex australis Erichson, 1842:270. TYPE: Lectotype 0, Tasmania,

Schayer [BERL, 5986; selection of Mattingly and Marks 1955:163 |.

1856. Culex crucians Walker, 1856:432. TYPE: *Holotype 9, Tasmania (Van

Diemen's Land) [BMNH; see Mattingly and Marks 1955:165]. Synonymy

with australis by Mattingly and Marks 1955:163, 166.

1911. Culex tasmaniensis Strickland, 1911:181. TYPE: *Lectotype & with gen-

italia mount, Tasmania, 21 Nov 1910, Bancroft (185) [BMNH: selection of

Belkin 1962: 329]. Synonymy with crucians by Edwards 1924:387.

1914. Caenocephalus concolor Taylor, 1914:700-701. TYPES: syntypes o and

¢, Cronulla, New South Wales, 13 Nov 1911, Mrs. J.B. Cleland [SYDN].

Synonymy with australis by Mattingly and Marks 1955:163.

1921. Aedes (Ochlerotatus) asl ashworthi Edwards, 1921:75-76. TYPE: *Holotype

o, Yallingup, West Australia, Sept 1914, J.H. Ashworth [BMNH]. Syn-

onymy with australis by Belkin 1962:329.

Aedes (Halaedes) australis of Belkin (1962:329-331); Stone (1963:128); Nye and

McGregor (1964:16); Dobrotworsky (1965:168-170; 1966:142).

Aedes (Pseudoskusea) australis of Mattingly and Marks (1955:163-166); Stone,

Knight and Starcke (1959); O'Gower (1959; 1960); Dumbleton (1962); Nye

(1962).

Aedes (Pseudoskusea) concolor of Edwards (1924:387; 1926b:113; 1932:158);

Taylor (1934:19); Woodhill (1936); Woodhill and Pasfield (1941: 207-208);

Lee (1944:75); Knight, Bohart and Bohart (1944:26, 51).

Aedes (Pseudoskusea) crucians of Edwards (1924:387: 1932:158); Taylor (1934:

19).

Aedes (Pseudoskusea) ashworthi of Edwards (1932:158); Taylor (1934:19); Stone,

Knight and Starcke (1959).

Belkin: Culicidae of New Zealand 95

As described and figured by Belkin (1962); in general similar to chatham-

icus; with the following diagnostic features, corrections and departure from its

other populations and from chathamicus.

FEMALE (Dobrotworsky 1965:fig.65). Wing: 5.5 mm. Proboscis: 3.4 mm.

Forefemur: 2.4mm. Abdomen: about 3.33 mm. Large dark species. Head:

Interocular space distinct but narrow, at its narrowest about 0.23 of its max-

imum height, without bristles or scales except at extreme upper end; frons a

little narrower than in chathamicus. Vertex with all the narrow decumbent

scales pale; erect scales shorter and broader than in chathamicus, more dis-

tinctly striated and with widened and forked apex. Labium largely dark-scaled,

ventral surface with scattered pale scales or indistinctly pale in basal 0.6 or

more. Palpus about 0.2 of proboscis; broader than in chathamicus, and with

distinctly smaller segment 5; segments 2 and 3 with small dingy pale scales,

a few similar pale scales at base of segment 4 particularly laterad, remainder

of segment 4 with dark scales. Antennae separated by a little less than diame-

ter of one torus; distinctly shorter than proboscis; torus and first flagellar

segment with numerous scales; bristles of flagellar whorls longer than in chat-

hamicus. Thorax: Mesonotum without linear light-scaled pattern; scales all

narrow, mainly bronzy but with scattered golden and light golden scales, more

numerous in supraalar area and prescutellar space. Mesonotal bristles much

less numerous and shorter than in chathamicus; fossals very small and few;

parascutellum without bristle. Scutellum with scales all narrow and predomi-

nantly pale; bristles less numerous than in chathamicus. Pleural scaling ex-

tensive, scales all broad, both dingy white and dark iridescent present; ssp

scale patch extensive, light; psp with numerous pale scales; ppn scales largely

dark; bristles less numerous and shorter than in chathamicus but 4,5 strong

anterior lower mep bristles present. Legs: Scaling dark except for usual light

areas of femora, very indistinct pale knee spots and apices of femora. Claws

with outer basal spicules and subbasal tooth, essentially as in chathamicus ex-

cept that hindclaws similar in size to others. Wing: Base of Sc without bris-

tles; wing scales all dark. Marginal scales of alula shorter and broader than

in chathamicus. Abdomen: Scaling of tergites II-VII largely dark, with varia-

ble complete basal transverse white or whitish bands usually narrowed middor-

sally and widened laterally; scaling of sternites II-VII largely pale, with apico-

lateral dark patches progressively larger on distal segments. Hairs on tergites

and sternites shorter than in chathamicus; sternite I without bristles. Female

Genitalia: essentially as in chathamicus but lobes of tergite IX much reduced.

MALE (Dobrotworsky 1965:fig.65; Mattingly and Marks 1955:fig.1). Es-

sentially as in the female and differing from chathamicus in the following. Pal-

pus about 0.8-0.9 of proboscis, dark-scaled; segment 5 swollen; segments 4

and 5 usually curled up in dry specimens; with numerous long bristles on ven-

tral and outer surface of apex of segment 3 and all of 4. Antenna markedly dif-

ferentiated; torus greatly enlarged; flagellar segments 1-11 short, with strong-

ly differentiated whorls of very long bristles on lateral and dorsal surfaces,

progressively shorter and fewer on distal segments, segment 1 with about 12

dorsal and 12 lateral, segment 13 with about 6 in each group; mesal whorl bris-

tles short but numerous; segments 12 and 13 greatly elongated, together sub-

equal to rest of flagellum, 13 a little longer than 12. Femoral and tibial light

spots more distinct than in female. Claws of all legs unequal; those of fore

and midlegs greatly enlarged, with anterior member of pair larger and witha

toothlike outer basal spicule and a submedian ventral tooth, posterior member

smaller and with numerous short subbasal spicules and a submedian ventral

96 Contrib; Amer. Ent. Inst), vol. 3, nov.4, 1968

tooth; anterior claw of hindleg slightly larger than posterior, both essentially

as infemale. Wing not shortened, abdomen shorter than wing.

MALE GENITALIA (Belkin 1962:fig.210; Nye and McGregor 1964:fig. 10;

Dobrotworsky 1965:fig.65; Mattingly and Marks 1955:fig.2). As figured by

Belkin; basically similar to chathamicus and with the following diagnostic fea-

tures. Basal tergomesal area of sidepiece with extensive patch of lanceolate

curved striated scales with narrow long base. Clasper slightly widened in bas-

al half. Claspette short, rounded, with numerous flattened curved hairs.

PUPA (Belkin 1962:fig.210). Abdomen: 3.98 mm. Trumpet: 0.54 mm.

Paddle: 1.24 mm. Trumpet short and broad, index about 2.2-2.4; moderately

flared; pinna distinctly oblique. Abdominal hair 1-I with more than 40 com-

pound branches; hair 5-II not closely associated with hair 4; hair 3-III cepha-

lad of hair 5; hair 5-IV, V usually double; hair 9-VII weak, removed consider-

able distance mesad from lateral margin of tergite. Paddle distinctly wider

than tergite VIII, paddle hair 1-P usually single.

LARVA (Belkin 1962:fig.211; Dobrotworsky 1965:fig.65; Lee 1944:fig. 52;

Woodhill and Pasfield 1941:fig.6). Head: 1.02 mm. Siphon: 1.09 mm. Anal

Saddle: 0.34 mm. Head hairs 5,6-C usually both single, weak; 13-C single,

weak. Antennal hair 1-A usually double (1-3). Thorax with mesothoracic

and metathoracic hairs close together and far removed from prothoracic. Pro-

thoracic hairs all weak and usually single or double. Metathoracic pleural group

(9-12-T) very small, dorsad of hair 8-M and separated from hairs 6,7-M by

little more than length of their common basal tubercle, hair 9-T small, usually

with less than 5 branches. Abdominal hairs weak; hair 6-VI about 0.5 of hair

4. Comb ina large patch of 100 or more short, broad, apically fringed scales.

Siphon characteristically shaped as in figure, index about 2.0-2.5; trachea

greatly expanded in siphon, black; acus not developed; pecten with 2 types of

teeth, those at base short and widely spaced, followed by 10-15 long teeth whose

bases are contiguous; hair 1-S at about middle of siphon, considerable distance

beyond pecten. Anal segment without any indication of acus; saddle restricted

to distal half of dorsal surface, hair 1-X far removed from it ventrad; ventral

brush usually with 7 pairs or 13 hairs all on grid.

SYSTEMATICS. The New Zealand population of australis does not appear

to differ in any significant feature from Australian populations I have examined.

Marks (in Nye 1962:33) suggested the possibility that it may have been intro-

duced from Australia and Nye (loc. cit.) pointed out that "there may be some

significance in the fact that these first records have come from the vicinity of

the ports of Bluff and Dunedin." I believe, however, that it is more likely that

australis reached New Zealand through natural dispersal rather than through

human agency. The discovery of chathamicus, which is undoubtedly a deriva-

tive from the same stock as australis, indicates that this phyletic line dispersed

to New Zealand in the past. I interpret the presence of australis as the result of

a more recent similar dispersal, perhaps contemporaneously with the disper-

sal of this species to Norfolk and Lord Howe islands. Such a dispersal may

have occurred through wind carriage of eggs from a relatively distant land

source to the east of New Zealand across a barrier too great for other species

of mosquitoes. The great tolerance of salinity shown by australis may have

been also a very important factor in its successful dispersal and establishment

in New Zealand.

As indicated under Opifex fuscus and A. chathamicus, australis is undoubt-

edly the most derived member of this common phyletic line. The similarities

with chathamicus particularly are very striking in all stages and in nearly all

Belkin: Culicidae of New Zealand 97

cases represent a derived state in australis, as follows: (1) in both sexes, nar-

rowing of the interocular space and frons; differentiation of flagellar segments

and bristlesin the flagellar whorls; decrease in bristles and increase in scales

on thorax and abdomen; complete loss of ventral bristles on vein Sc; (2) differ-

entiation of the antenna, palpus and claws in the male; (3) reduction of tergite

IX in the female genitalia; (4) development of specialized scales on the side-

piece and hairs on the claspette of the male genitalia; (5) in the pupa, displace-

ment of hair 3-II and displacement of hair 9-VII; and (6) in the larva, reduction

and greater approximation of metathoracic pleural to mesothoracic hairs; re-

duction in size of hair 6-VI, pecten, anal saddle and ventral brush; loss of acus

in siphon and anal segment.

BIONOMICS. The breeding sites of australis in New Zealand appear to be

Similar to those occupied by O. fuscus, are within splashing or seepage dis-

tance of the ocean, and show salinities ranging from 0.3 to 3.6% NaCl (Nye

and McGregor 1964:16). In Australia this species breeds only in salt or brack-

ish rock and ground pools at or just above high-tide mark (Mackerras 1926;

Woodhill 1936:634; Woodhill and Pasfield 1941:208; Dobrotworsky 1965:170).

Experimental studies by Woodhill (1936) indicate that the immature stages have

extreme tolerance of gradual concentration of sea-water (emergence of some

adults in sea-water with 18 G NaCl and survival of larvae for 5 days in 20%

NaCl). O'Gower (1959; 1960) has reported on the oviposition behavior and lar-

val ecology of australis in the Sydney area.

In Australia this species is considered to be animportant pest in restricted

localities a few hundred feet from the breeding sites near the coast (Woodhill

1936:633-634) but australis has not been reported to date to bite man in New

Zealand. Experimental studies by Woodhill (1936:634) have shown autogeny

and stenogamy in australis.

DISTRIBUTION (fig. 38). Material examined: 1957 specimens; 15 0, 41 9

1713 larvae, 188 pupae; 35 individual rearings (22 larval, 11 pupal, 2 incom-

plete). :

AUSTRALIA. Widely distributed along the southeast coast from southern

Queensland, through New South Wales, Victoria, Tasmania, South Australia to

southern part of Western Australia. LORD HOWE ISLAND. NORFOLK ISLAND.

NEW ZEALAND. Westland: Whataroa, o, M. Pillai [Nye and McGregor

1964: 16].

Otago: Shag Pt. and Karitane [Nye and McGregor 1964:16]. Otago Penin-

sula, 19 Aug 1962, E.R. Nye, 1% [UCLA]; Boulder Beach, 28 Apr 1962, E.R.

Nye, 1¢ [UCLA]; Hooper's Inlet, 2 Jan 1964, BN (NZ 70), 9 1p? (70-101-

103, 105-110), 1 p? (70-104), 77, 22, 187 L, 10 P[UCLA]; Papanui Inlet, 19

Aug 1962, E.R. Nye, 1c’ [UCLA]; Cape Saunders, Papanui Inlet, Penguin

Beach, Pipikaretu Beach, Potato Point and Purakanui [Nye and McGregor 1964:

16].

Southland: Bluff, 26 Jan 1964, S (NZ 104), 13 1p? (104-101-107, 185-190),

9 p? (104-108, 109, 114-120), 2 1p (104-110, 191), 5, 69, ca 1500 L, 143 P

[UCLA]; Bluff, D.D. McGregor [Nye and McGregor 1964:16].

Stewart Island: Deep Bay, boat hull, 21 Dec 1961 [Nye and McGregor]; 27

Jan 1964, S(NZ 106), 1 po’ (106-105), 2 L [UCLA].

Subgenus Ochlerotatus

1891. Ochlerotatus Lynch Arribalzaga 1891a:374; 1891b:143. TYPE SPECIES:

98 : Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

O. confirmatus Lynch Arribalzaga, 1891, Argentina; selection of Co-

quillett (1910:577).

For complete synonymy see Stone, Knight and Starcke (1959).

20. Aedes (Ochlerotatus) antipodeus (Edwards)

Fig. 39

1920. Ochlerotatus antipodeus Edwards, 1920:132. TYPE: *Holotype “, Kari-

kari Bay, New Zealand, 5 Feb 1916, A. E. Brookes [BMNH; marked as

_ type by Edwards ].

Aedes (Ochlerotatus) antipodeus of Edwards (1924:376; 1932:137); Taylor (1934:

15); Graham (1939:212-213); Miller (1950:42); Miller and Phillipps (1952:

16-17); Stone, Knight and Starcke (1959); Belkin (1962:389-391); Marks and

Nye (1863:50-54); Nye and McGregor (1964:17-20).

As described and figured by Belkin (1962:389-391) and Marks and Nye

(1963:50-54); with the following diagnostic features.

FEMALE (Nye and McGregor 1964:fig.12; Marks and Nye 1963:fig. 1;

Miller and Phillipps 1952:figs. 28-30. Wing: 5.16 mm. Proboscis: 3.33 mm.

Forefemur: 2.25 mm. Abdomen: about 3.5mm. Medium to large dark spe-

cies. Vertex of head with a pair of large submedian patches of short narrow

dark decumbent scales; palpus and proboscis entirely dark. Mesonotum pre-

dominantly with narrow dark bronzy scales and with a more or less conspicu-

ous and contrasting pattern of golden scales in narrow lines and patches; pre-

scutal golden line incomplete, ending before scutal angle; parascutellum with-

out bristles. Scutellum with broad median line of narrow golden scales, re-

mainder of scales narrow and dark. Posterior pronotum predominantly with

flat broad dark scales; ssp usually without scales; psp usually (always?) with

a few whitish broad scales; lower msp bristles absent. Femora not mottled;

knee spots inconspicuous; tarsi with narrow basal whitish bands, foretarsus on

segments 1 and 2 (Sometimes also 3 and 4), midtarsus on 1-3 (Sometimes also

4), hindtarsi on 1-4 (sometimes also 5). Claws all with fine spicules on outer

surface at base; those of foreleg and midleg with a subbasal inner tooth. Wing

scales all dark; vein Sc without basal bristles ventrally. Abdominal tergites

II- VII with narrow to moderately broad basal transverse whitish bands connec-

ted to larger basal lateral white patches; sternites dark-scaled, with basal lat-

eral white patches. |

MALE (Marks and Nye 1963:fig.1). Essentially as in the female except for

sexual differences. Palpus variable in length, usually exceeding proboscis

by about length of segment 5, sometimes slightly shorter than proboscis; seg-

ments 4 and 5 down-turned and with basal pale scale patches. Fore and mid-

tarsal claws unequal; enlarged clawof foretarsus with a basal and a submedian

tooth, that of midtarsus with basal tooth only.

MALE GENITALIA (Belkin 1962:fig. 264; Marks and Nye 1963:fig.1; Nye

and McGregor 1964:fig.13). As figured most accurately by Marks and Nye.

Basal tergomesal lobe strongly developed, with numerous short setae on tergal

surface, those on mesal margin of lobe longer and slightly thickened. Clasper

strongly swollen in the middle. Claspette filament bristlelike, subequal in

length to body of claspette. |

PUPA (Belkin 1962:fig.264; Marks and Nye 1963:fig.2). Abdomen: 3.4

mm. Trumpet: 0.54 mm. Paddle: 0.94 mm. Chaetotaxy as figured by Belkin

Belkin: Culicidae of New Zealand 99

and as figured and described by Marks and Nye. Trumpet uniformly darkly

pigmented except sometimes for a more strongly darkened apex; usually dis-

tinctly narrowed before apex; with slightly oblique pinna. Abdominal hair 2

Sspiniform on all segments, submedian in position on HI-VIJ; hairs 6-1, II, 7-I,

3-III and at least 5-IV, V all very long, thickened and usually all single.

LARVA (Belkin 1962:fig.265; Marks and Nye 1963:fig.2; Nye and McGre-

gor 1964:fig.14. Miller and Phillipps 1952:fig.31). Head: 0.96 mm. Siphon:

1.37 mm. Anal Saddle: 0.39 mm. Chaetotaxy as figured by Belkin and described

by Marks and Nye. Head hair 1-C moderately heavy and about as long as dis-

tance between bases; 6-C usually single; 5-C usually at least triple. Protho-

racic hairs 2, 3-P usually branched, weak; 4-P multiple; 8-P long and usually

single. Abdominal hairs 1-IV, V weak, usually at least triple. Comb scales in

a large patch of 30-45; primarily spatulate and fringed apically; outer scales,

particularly ventrally, spinelike and with fringe at base only. Siphon index 3.0-

4.0; pecten teeth 17-22, very darkly pigmented, evenly and closely spaced

throughout, progressively longer distad; hair 1-S distad of pecten, at about

middle of siphon. Anal segment with poorly pigmented detached acus ventrally

as figured by Belkin; ventral brush with at least 7 pairs of hairs on grid, usu-

ally with 1 additional, unpaired hair, and usually with 1 precratal hair, some-

times with 2-4.

SYSTEMATICS. A. antipodeus has been placed in the Vigilax Section of

Ochlerotatus by Marks (1957:72) on the basis of features of the male genitalia

and the larval siphon and comb. It appears to be related to inexpectatus Bonne-

Wepster, 1948 from the highlands of New Guinea, which it closely resembles in

scale pattern of the adults, in the male genitalia and pupa but not in larval fea-

tures (Marks and Nye 1963:54).

There is considerable variation in the morphology of all stages of antipodeus

but no indication in the material at hand of any geographical differentiation.

BIONOMICS. A. antipodeus is evidently a typical flood water ground pool

breeder. Immature stages have been found in temporary pools, margins of

flooded streams, reservoirs and swamps, and in stream bed pools. All breed-

ing sites for which some data is available have been in partial or dense shade

in native bush. Breeding occurs primarily during the colder months of the

year, from March through October, but males have been found in November,

December, January and February suggesting that at least limited breeding takes

place during the summer months. Miller and Phillipps (1952:17) report that

F.E. Ellett found larvae quite active during the day in a pool whose nocturnal

temperature fell to 13°F and Graham (1939:213) found full development to the

pupal stage at temperatures no higher than 420F and completion of the entire

aquatic cycle in 3 weeks at 52°F. Females have been observed laying eggs on

slime and mud in the winter and larvae have been found in abundance within 24

hours of flooding of dry depressions (Graham, loc. cit.)

Females are active throughout the year and bite at any time’of day or night

(Graham 1939:212-213). Several have been collected in buildings but there are

no definite records of their biting humans indoors. Judging by the number of

collections at hand antipodeus is readily attracted to humans out-of-doors but

the small number of individuals represented in most collections suggests that

populations of this species are seldom large. There are no reports of antipo-

deus attacking cattle or other animals.

DISTRIBUTION (fig. 39). Material examined: 194 specimens; 20 o, 96 9,

46 larvae, 32 pupae; 5 individual rearings (3 larval, 2 incomplete).

NEW ZEALAND. North Auckland: Karikari Bay, 5 Feb 1916, A. E. Brookes,

100 Contrib. Amer. Ent. Inst., vol. 3, no..1, 1968

holotype ¢, 13¢ [BMNH]. Kaitaia, 5 Feb 1917, D. Miller, 10, 19 [BMNH],

1 o& [USNM]. Whangaroa, 25 Jan 1917, D. Miller, 19 |[BMNH]. Russell, 24

Jan 1917, D. Miller, 20, 19[BMNH], 19[USNM]. Waipoua Forest, 5-7 July

1937, J.M. Valentine, 1? [BISH]; Waipoua, D.H. Graham, 1% [Marks and

Nye 1963:52]. Nihotupu, 25 Feb 1923, A.L. Tonnoir, 19 [NELS]. Waitakeres,

20,26 Feb 1950, M. Knight, 22 [AUCK]; in house, 27 Dec 1950, K.P. Lamb,

1 [AUCK]. Auckland Watershed Reservoir, 26 Feb 1944, G. Chamberlain,

99 [WELL]. Titirangi, 1 July 1946, G.L. Bissett, 1 L (972), 8 L (973), 7L

(974) [NELS]; D.H. Graham, 20 Jan 1927,1 L, 11 Sept 1929, 1 L, 13 Sept 1929,

6 P [Marks and Nye 1963:52,54]. Auckland, Herne Bay, 24 Feb 1918, T.R.

Harris, 19 [BMNH]. |

Little Barrier Island: Small stream between Te Waikohare and Tirikikawa

streams, 9 Oct 1963, S(NZ 11), 19, 3 L, 3 P[UCLA]; 6 Mar 1964, S (NZ 128),

2L [UCLA]. Summit Track, elev. 267 m, 9 Oct 1963, S(NZ 12), 1 lpo& (12-

107); Bly (12> 106; 108). 1 p (12-102, 109), 4078-9, 12 L, 17 P{UCLAJ. In-

Side bunkhouse, 29 Oct 1963, S(NZ 18), 2 [UCLA]. Ranger's house, 30 Oct

1963, S(NZ 19), 52 [UCLA]. Locality unknown, 7 Nov 1963, S (NZ 33), 19

[UCLA ]; 25 Nov 1954, R.A. Harrison, 2 9 [AUCK]; 4 Dec 1929, 1 o%, 2 9 [AUCK].

Gisborne: Lake Waikaremoana, 8 ¢ |Marks and Nye 1963:51 |.

Wellington: Mt. Ruapehu, Jan 1921, M.N. Watt, 12 [USNM]. Ohakune, 15

July 1922-15 Jan 1923, T.R. Harris, 1o% [BMNH]. Palmerston North, 11 Aug

1944, Reid (399), 1 L [NELS]. Tehoro, 16 Feb 1911, R.F.L. Burton, 3 9

[BMNH]. Paraparaumu, Jan 1925, 19 [WELL]. Masterton, 15 Aug 1944, F.E.

Ellett (398), 4.1L, 5 P [NELS]; 28 Aug 1944 (416), 2 L [NELS]; 15 Aug 1945,

W.J. Phillipps (795), 1%, 39 [NELS]. Day's Bay, 29 Nov 1921, A. L. Tonnoir,

1 o [NELS].

Nelson: Canaan Beech Forest, 4 Jan 1964, BG (NZ 76A), 12 [UCLA].

Tasman, 10 Oct 1922, R.J. Tillyard, 19[NELS]. Nelson, 28 Sept 1923, A. L.

Tonnoir, 12 [NELS]. Tahunanui, 10 Mar 1964, S (NZ 157), 10%, 1L, 2 P

[UCLA]. Aniseed Valley, 8 Sept 1922, Philpott, 1 o [CANT].

Westland: Lake Moana [Brunner], 16-21 Dec 1925, A. L. Tonnoir, 30, 8°

[CANT].

Canterbury: Kaiapoi, 21 Jan 1964, S (NZ 99), 79[UCLA]. New Brighton,

11 May 1921, 1 9 [BMNH]; 5 Nov 1921, 1¢ [CANT]; Dyer's Road, 11 Nov 1922,

J.W. Campbell, 10, 12 [BMNH], 1% [CANT]. Christchurch, 21 Oct 1924,

A.L. Tonnoir, 10%, 32 [CANT]. Akaroa, 12 Dec 1924, A.L. Tonnoir, 1 9

[CANT]. Mt. Grey, Jan 1925, J.W. Campbell, 1 9 [BMNH].

Otago: Otago Peninsula, Papanui Inlet, Pipikaretu Beach, Taiaroa Head

and near Victory Beach [Nye and McGregor 1964:20]. Taieri Plain, near Dun-

edin [Nye and McGregor 1964:20]. Stuart's Gully, 6 Nov 1921, C. P. Alexander,

3.0, 49 [USNM].

Stewart Island: Locality not specified, coll. Hutton, 1 9 [CANT].

21. Aedes (Ochlerotatus) subalbirostris Klein and Marks

Figs. 22, 23, 39

1960. Aedes (Ochlerotatus) subalbirostris Klein and Marks 1960:115-116.

TYPE: *Holotype 9, Invercargill, New Zealand, 23 Nov 1886, W. Wesché

| BMNH |.

Aedes (Ochlerotatus) subalbirostris of Stone (1961:39); Belkin (1962:395-396);

Marks and Nye (1963:56-60); Nye and McGregor (1964:20-22).

Belkin: Culicidae of New Zealand 101

Aedes (Ochlerotatus) albirostris of Edwards (1925:376; 1932:137); Taylor (1934:

17); Miller (1950:42); Stone, Knight and Starcke (1959).

As described by Klein and Marks (1960:115-116) and as described more

fully and illustrated by Nye and Marks (1963:56-60); with the following diagnos-

tic features.

FEMALE. Wing: 5.5mm. Proboscis: 3.0 mm. Forefemur: 2.25 mm.

Abdomen: about 3.5mm. Large dark species. Vertex of head without distinct

submedian patches of dark scales; palpus predominantly dark-scaled but with

some whitish scales on apices of segments and scattered elsewhere; proboscis

with variable amount of pale-scaling usually restricted to basal 0.75 (except

for extreme base), sometimes forming a more or less distinct broad subme-

dian pale ring. Mesonotum without distinct linear pattern, mainly with narrow

golden scales except for patch of creamy scales above wing root and prescu-

tellar area; parascutellum without bristles. Scutellum with all scales narrow

and creamy. Posterior pronotum with narrow curved creamy scales above, a

variable number of elongate flat black scales in the middle and broad creamy

white scales below; ssp with a large patch of broad whitish scales, sometimes

divided; psp with similar scales; lower msp bristles absent. Femora mottled,

predominantly pale-scaled; knee spots inconspicuous; tibiae with some pale

scales at base and apex; tarsi with very narrow basal whitish bands, foretar-

sus on segments 1-3 and sometimes some white scales on 4, midtarsus on seg-

ments 1-3 and some basal white scales on 4 and 5, hindtarsus on segments 1-4

and a patch on 5. Claws all with fine spicules on outer surface and all with sub-

basal inner tooth. Wing with dark scales except for small patch of pale scales

at base of vein C. Abdominal tergites I-VII with more or less complete straight

narrow basal pale transverse bands confluent with basal lateral triangular white

patches, some pale scales apically and sometimes near middle of tergites; ster-

nites predominantly pale-scaled, with some dark scales near base or lateral

preapical dark patches which may be joined.

MALE (Marks and Nye 1963:fig.2). Essentially as in the female except

for following sexual differences. Palpus exceeding proboscis by about 0.33 of

segment 5; apparently up-turned from base of segment 3; scales dark except

for pale band at base of segment 3 and base of 4 and sometimes at few pale

scales at base of 5; numerous long hairs on apex of segment 3 and on segments

4 and 5. Labium entirely dark-scaled or with a few slightly paler scales

ventrally. Femora darker than in female; tarsi with pale scaling even more

restricted. Fore and midtarsal claws unequal, enlarged claw on both legs with

a basal and a submedian tooth. Wing without pale scales at base. Abdominal

tergites with straight basal transverse pale bands, no apical light scales on II-

V; sternites largely dark-scaled except for basal lateral pale patches and usu-

ally apical median pale patches.

MALE GENITALIA (fig.22; Marks and Nye 1963:fig.3; Nye and McGre-

gor 1964:fig.15). As figured here and as described and figured by Marks and

Nye; with the following diagnostic features. Sidepiece with prominent apical

tergal lobe with minute setae on its mesal and sternal surface and 2 conspicu-

ous long differentiated bristles nearits base on tergal surface; basal tergomes-

al lobe not prominent, largely mesal in position, with numerous moderately

long setae, none differentiated; sternal surface toward the mesal margin with

3,4 rows of mesally directed long setae, the most lateral of these very heavy,

others thin. Clasper slightly expanded in basal half. Claspette filament almost

1.5 of body, flattened and slightly expanded in distal 0.67 before the curved

102 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

tapered tip.

PUPA (fig.22; Marks and Nye 1963:fig.3). Abdomen: 4.25 mm. Trum-

pet: 0.77 mm. Paddle: 1.2 mm. Chaetotaxy as figured here and as described

by Marks and Nye. Trumpet uniformly moderately pigmented except for slight-

ly darkened apex; distinctly flared apically; pinna distinctly oblique. Abdomi-

nal hair 2 thin on all segments, in distal 0.25 on IJI-VII; hairs 6-I, II, 7-I all

long, single, moderately thickened except for thin attenuate apex; 3-III, 5-IV-

VI moderately long, moderately thickened and usually all double.

LARVA (fig.23; Marks and Nye 1963:fig.4; Nye and McGregor 1964:fig.

16). Head: 1.08 mm. Siphon: 1.45 mm. Anal Saddle: 0.45 mm. Chaetotaxy

as figured here and as described by Marks and Nye. Head hair 1-C thin and

distinctly shorter than distance between bases; 5, 6-C both usually double. Pro-

thoracic hairs 2, 3-P both single and thickened; 4-P single or double; 8-P very

short, branched. Abdominal hair 1-IV,V thickened, usually double, rarely

triple. Comb scales in a very large patch of 60-100 in about 5 rows; scales

small with widened rounded tip, finely fringed throughout. Siphon index about

2.6-3.5; pecten teeth 13-17, very darkly pigmented, 1-3 distal teeth widely

separated from others and considerably larger; hair 1-S distad of pecten, at

about 0.67 from base of siphon. Anal segment with very poorly pigmented de-

tached elongate acus ventrad of saddle as figured; ventral brush usually with

at least 7 pairs of hairs on grid (8 according to Marks and Nye) and 2-3 pre-

cratal unpaired hairs.

SYSTEMATICS. This species was confused with albirostris (Macquart,

1850), whose type locality was erroneously indicated as Akaroa by its author,

until Klein and Marks reexamined the type material of the latter and the single

femalefrom Invercargill which Edwards provisionally referred to as albirostris

despite a number of discrepancies with the original description. Klein and

Marks showed that albirostris came from New South Wales and that it is con-

specific with vigilax (Skuse, 1889), and described subalbirostris as a new spe-

cies on the basis of the unique specimen from Invercargill. Additional females

were captured by L.J. Dumbleton (1963:26-27) in the Catlins district in 1958

and 1959 but the immature stages and the males were not known until 1962 when

E.R. Nye and D. McGregor found the breeding sites of this species on the Ota-

go Peninsula and associated all the stages through rearings.

A. subalbirostris shows sexual dimorphism in leg markings, an unusual

condition in Australian Ochlerotatus according to Marks and Nye (1963:59). In

the opinion of these authors it appears to fall into the Perkinsi Section as de-

fined by Marks (1957) and modified by Dobrotworsky (1960) but seems to be no

more closely related to the Australian members of this section than to 2 South

African species which are allied to this section (Marks 1959). Marks and Nye

(1963) consider that the present knowledge of this ancient group is insufficient

to suggest whether subalbirostris represents a derived or a primitive stock.

BIONOMICS. A. subalbirostris has been found breeding in all but one in-

stance in association with antipodeus in ground pools where it was the domi-

nant species. It has been recorded once breeding alone in a horse trough on

the Taieri Plain. In Otago both species favor clean fresh water even in close

proximity to the sea (Nye and McGregor 1963:20).

Females of subalbirostris are attracted to man occasionally and may enter

buildings as indicated by their capture in the Catlins District (Dumbleton 1963)

and in a hotel (holotype). As in the case of antipodeus, populations of subalbi-

rostris apparently do not reach high levels of density.

DISTRIBUTION (fig.39). Material examined: 27 specimens; 5c, 69, 11!

Belkin: Culicidae of New Zealand 103

larvae, 5 pupae; 5 individual larval rearings (3 co, 2 9).

NEW ZEALAND. Otago: Otago Peninsula, Hooper's Inlet, 3 Aug 1963,

N (NZ 223), 3 lpo& (223-10, 11, 13), 2 lp? (223-12, 14), 6 L [UCLA]; Hooper's

Inlet, Papanui Inlet, Pipikaretu Beach [Marks and Nye 1963:59]; Tairoa Head

and near Victory Beach [Nye and McGregor 1964:20]. Taeri Plain, near Henley,

23 Sept 1962, E.R. Nye, 2%, 32 [UCLA]; Taeri Plain [Marks and Nye 1963:

59]. Catlins District [Marks and Nye 1963:56].

Southland: Invercargill, 23 Nov 1906, W. Wesche, holotype ? [BMNH].

Subgenus Finlaya

1903. Finlaya Theobald, 1903:281-283. TYPE SPECIES: Culex kochi Doenitz,

1901, New Guinea; selection of Blanchard (1905:415).

For complete synonymy see Stone, Knight and Starcke (1959).

22. Aedes (Finlaya) notoscriptus (Skuse)

Fig. 40

1889. Culex notoscriptus Skuse, 1889:1738-1740. TYPES: Syntypes, o, 2, Syd-

ney, Australia, Masters and Skuse (MACL].

Aedes (Finlaya) notoscriptus of Edwards (1922:100; 1924:382; 1932:152); Gra-

ham (1929:205-215); Taylor (1934:18); Woodhill and Pasfield (1941:205);

Lee (1944:57); Knight, Bohart and Bohart (1944:38, 53); Knight and Marks

(1952:558-559); Laird (1954:286-287); Iyengar (1955:27; 1960:61); Rageau

(1958a:877; 1958b:3); Stone, Knight and Starcke (1959); Belkin (1962); Dob--

rotworsky (1965:131-133; 1966:140-141); Steffan (1966:208).

Aedes notoscriptus of Graham (1939:212); Perry (1950:112); Miller (1950:42-

43); Miller and Phillipps (1952:12-15).

Ochlerotatus notoscriptus of Miller (1920).

As described and figured by Belkin (1962:348-349); with the following diag-

nostic features and significant differences from extralimital populations.

FEMALE (Dobrotworsky 1965:fig.49; Miller and Phillipps 1952:figs. 19,

21-23,25; Graham 1929:figs.1-10). Wing: 4.1 mm. Proboscis: 2.6 mm.

Forefemur: 2.15 mm. Abdomen: about 3.0 mm. A small to medium-sized

brown mosquito with conspicuous narrow predominantly golden lines forming a

lyre-shaped pattern on mesonotum, scutellum with broad silvery scales, pro-

boscis and tarsi banded with white. Head: Orbital line silvery; median, pos-

terior and lateral decumbent scales of vertex golden, remainder blackish; pro-

boscis with narrow submedian ring of pure white scales; palpus, tipped with

Silvery scales. Thorax: Acrostichal and dorsocentral bristles strongly devel-

oped, in complete rows; integument light brown. Acrostichal light-scaled line

and its prescutellar forks silvery with slight yellowish tinge; anterior dorso-

central light line always present and long, almost reaching anterior border,

distinctly golden; posterior fossal line broadened caudally, distinctly golden;

lateral prescutal-posterior fossal-posterior dorsocentral light line continuous,

broadened from scutal angle to prescutellar space, usually silvery except from

posterior end of fossal line to prescutellar space where it is golden; supraalar

line long, silvery; prealar patch silvery; silvery broad scales of scutellum

104 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

with slight yellowish tinge; paratergite and pleural scales patches silvery ex-

cept on upper part of ppn where they are blackish. Legs: Longitudinal light-

scaled lines of femora and tibiae dingy white; fore and midtarsi with small bas-

al white markings only on segments I and 2; hindtarsal white markings usually

present on segments 1-4 only, narrow, about 0.25 of segment length on 2, seg-

ment 5 light brown, Sometimes with a few to numerous white scales at base

dorsally. Abdomen: Tergites usually with distinct median dorsal basal white

or dingy markings widened in the middle but not joined to lateral silvery spots;

predominant scaling of sternites varied from light golden to dark.

MALE (Miller and Phillipps 1952:fig.24; Graham 1929:fig.11). Essen-

tially as in the female except for usual sexual differences; lateral light line of

mesonotum usually not as distinctly broadened as in female and usually silvery

throughout; hindtarsal segment 5 usually with more numerous white scales;

abdominal tergites IJJ-V usually with broad straight basal dorsal bands joined

to lateral silvery spots.

MALE GENITALIA (Dobrotworsky 1965:fig.49; Belkin 1962:fig.226; Gra-

ham 1929:fig.12). Apparently indistinguishable from other populations. Side-

piece long and slender; lanceolate-tipped setae varied in number and extent.

Clasper shorter and more swollen in basal 0.5 than shown by Belkin (1962) and

Dobrotworsky (1965).

PUPA (Belkin 1962:fig. 226; Graham 1929:figs. 23,24). Abdomen: 4.0 mm.

Trumpet: 0.6 mm. Paddle: 1.0mm. Diagnostic characters as in the key;

chaetotaxy as figured by Belkin (1962) for New Caledonia population, except as

noted and with considerable variation in branching. Abdominal hair 1-II with

3-10 branches, usually simple, rarely weakly dendritic; hairs 3-III, 5-IV, V

weaker and shorter, especially 3-IIIJ, none usually as long as tergite following.

LARVA (Belkin 1962:fig.227; Dobrotworsky 1965:fig.49; Miller and Phil-

lipps 1952:fig. 26; Lee 1944:fig.35; Woodhill and Pasfield 1941:fig.3; Graham

1929:figs. 14-22). Head: 0.89 mm. Siphon: 0.55 mm. Anal Saddle: 0.32 mm.

As figured by Belkin (1962); diagnostic characters as in the key. Larger than

in most other populations; comb scales in middle of posterior row elongate;

comb scale patch variable; siphon index about 2.0 or slightly less; saddle hair

usually with 3 or more branches.

SYSTEMATICS. The adults of the New Zealand populations of notoscriptus

have a very characteristic facies quite different from that of any of the other

population I have seen. They are characterized by the following features in the

female: (1) a considerably larger size, (2) predominantly golden lines on the

mesonotum, especially the long anterior dorsocentral one, (3) hindtarsal seg-

ment 5 largely dark, and (4) basal abdominal light bands widened in the middle

and not reaching the lateral silver spots. However, all of these features ex-

cept the larger size are found sporadically in extra-limital populations of noto-

scriptus which exhibits a very wide range of variation of coloration throughout

its extensive known range. Similarly the characteristic, rather constant, fea-

tures of the immature stages mentioned in the diagnosis above occur sporadic-

ally in other populations, notably in some populations in Australia and New Cal-

edonia. The male genitalia of the New Zealand populations show no significant

differences from other populations. Therefore, it appears that the New Zea-

land populations are nothing more than a segregate of notoscriptus whose lar-

ger size and color differences may be associated with the colder environment

in New Zealand.

Graham (1939:212) reported notoscriptus "having been more than once

taken alive on vessels on their arrival at Auckland from Sydney" and, on this

Belkin: Culicidae of New Zealand 105

basis and the apparent occurrence of this species only around Auckland, Nel-

son and Whangarei at that time, considered it to be a recent introduction to

New Zealand. I have not been able to locate any of the specimens recovered

from ships but there is no reason to doubt that notoscriptus was the species

reported by Graham. Although at the present time the known distribution of

notoscriptus in New Zealand is greater than in Graham's time it is still entire-

ly restricted to the vicinity of present day or former seaports. Unlike Culex

quinquefasciatus, the other introduction to New Zealand, notoscriptus is not

confined to man-made habitats and has invaded nearly all the other habitats a-

vailable in New Zealand similar to those it has been reported to utilize else-

where. Although this circumstantial evidence is not unequivocal, it is highly

suggestive that notoscriptus is indeed an introduced species in New Zealand.

BIONOMICS. During our survey, the New Zealand population of notoscrip-

tus was found to breed largely in various types of large and small artificial con-

tainers, wood, metal and cement, but it was also recorded from pools in dry-

ing stream beds (NZ 128,130), rot holes in native trees (NZ 37, 38, 47,92) and

astelia axils (NZ 60,67). Graham (1929:210) records it in addition from rock-

holes, fronds of nikau palms (on the palm and on the ground), in banana axils

and occasionally in drains, swamps and waterholes. Densely shaded habitats

appear to be preferred by this species.

A. notoscriptus is a diurnal biter. It attacks man readily but according to

Graham (1929:210) it prefers cattle as a source of blood. Apparently biting

activity occurs only in the shade. Graham reports it as a frequent intruder in

houses in Auckland but in general it is a sylvan species and is especially abun-

dant in densely wooded gullies. According to Graham (1939:212) notoscriptus

passes the winter in both adult and larval stages.

DISTRIBUTION (fig. 40). Material examined: 2882 specimens; 126 o, 171

?, 2176 larvae, 409 pupae; 109 individual rearings (67 larval, 37 pupal, 5 in-

complete).

Widely distributed in AUSTRALIA, NEW CALEDONIA, LOYALTY ISLANDS,

NEW BRITAIN, NEW IRELAND, NEW GUINEA, and MOLUCCAS.

NEW ZEALAND. North Auckland: Opua, 20 Dec 1963, BS (NZ 37, 38),

6 lpo (37-102, 104,105; 38-101, 102,104), 51p? (37-103, 106-108; 38-103),

1d p(37-101), 22 (37) 1 9 (88),°10 17182) 6: (38),1. B87), 2 P88) PL GCLA).

Whangarei (Graham 1939:212). Dargaville, 12 Dec 1944, G.L. Bissett (494),

1 L[NELS]. Warkworth, 29 Nov 1944, G.L. Bissett (479), 1 L[NELS]. When-

uapai, 31 Oct 1944, G.L. Bissett (441), 1 L [NELS]; 15 Jan 1945, G.L. Bis-

sett (534), 1 L[NELS]. Brown's Bay, 18 Feb 1945, H. Harrison, 1 2? [AUCK].

Takapuna, 12 May 1944, A. Sanford (319), 1 L[NELS]. Rangitoto Island, Sum-

mit Track, 14 Sept 1963, S (NZ 6), 3 lpo (6-101, 109, 111), 1 lp? (6-107), 1 po

(6-103); 2 p2 (6-102; 106); 2 1p (6-108, 110), 39, 13 L, 3 P (UCLA) gun em-

placement, 14 Sept 1963, S (NZ 8), 21po (8-109, 112), 4 1p? (8-110, 111, 113,

114), 1 po (8-101), 4 p? (8-102-105), 2o, 29, 12 L, 9 P [UCLA]; lava flow

near sea, 4 Jan 1964, S (NZ 67), 1 po (67-112) [UCLA]. Henderson, 8 Nov

1944, G.L. Bissett (458), 1 L[NELS]. Waitakere Ranges, Quarry tributary of

Waitakere River, 28 Dec 1963, BF (NZ 59,60), 1 lpo& (60-112), 12 (59)

[UCLA]. Bethells Road Swamp, 10 Jan 1965, F (NZ 200A), 19[UCLA]. Titi-

rangi, S. Titirangi Road, 11 Jan 1964, BS (NZ 95), 21lpo (95-102,108), 5

1p? (95-101, 104-107), 2 po (95-109, 110), 1 1p (95-103), 60°, 592, 68 L, 20 P

[UCLA]. Titirangi Beach, end of Mahoe Road, 30 Dec 1963, BF (NZ 614,

61B, 63), 21lpo (63-104, 105), 12 lp? (61B-101, 108-114, 116; 63-101-103),

Llp (61B+115), 8:4..(61)) :20:¢ .(63),: 27 2 (61); 2:9 (63), 706: (61), 43 1:(63),

106 Contrib. Amer. Ent. Inst.,°vol. 3, no. 1, 1968

35 P (61), 14 P (63) [UCLA]; 11 Jan 1964, BS (NZ 89), 21 2 [UCLA]; 11 Jan

1964, BS (NZ 91), 51p? (91-101-105), 38 Oo, 20 2, 3396 L, 85 P|UCLA|; 11

Jan 1964, BS (NZ 92), 1 lp? (92-120) [UCLA]; Mahoe Road, 4 July 1964, F,

1 o (NZ 191A), 3 L (NZ 191B) [UCLA]. Laingholm, 10 Jan 1946, Goi: Bissett

(833), 21L[NELS]. Auckland, 1 L (455), 9 Jan 1945, G.L. Bissett (518), 1 L

[NELS]; Avondale, June 1950, H. Harrison, 3 9 [AUCK]; Methuen Road, 9 Aug

1964, F (NZ 194), 31lpo’ (194-126-128), 2 po’ (194-125, 130), 1 p? (194-129)

[UCLA]; Grafton, 18 July 1918, D. Miller, 10%, 19 [BMNH]; Greenlane, 6

Sept 1948, K.P. Lamb, 1 9 [AUCK]; Mt. Eden, 16 Apr 1944, E.M. Armstrong

(248), 1 L [NELS]; Remuera, 4 Feb 1946, G. L. Bissett (869), 1 L [NELS].

Mangere, 3 Feb 1950, K.P. Lamb, 19 [AUCK]. Localities uncertain Brough-

ton Rd, 24 Mar 1919, D. Miller, 19 [BMNH]; St. Mary's Bay, 24 Mar 1919,

D. Miller, 1%, 22 [BMNH]; Glenalvon, 20 Mar 1919, D. Miller, "in banana

palm," 2 ° [BMNH].

Little Barrier Island: Small stream between Te Waikahere and Tirikikawa

streams, 6 Mar 1964, S (NZ 128), 41po (128-106, 108, 110,121), 4 1p? (128-

114, 116, 125, 126), 6 po’ (128-107, 109, 111, 113, 119, 123), 7 p@ (128-103-105,

112,117, 122,124), 7%, 299, 155 L, 70 P[UCLA]. Near chicken house, 6 Mar

1964, S (NZ 129), 1 1p? (129-106), 4 po’ (129-115, 119, 121, 123), 5 p? (129-

101, 105, 107, 118, 120), 50°, 69, 39L, 36 P[UCLA]. Waipawa stream, 6 Mar

1964, S (NZ 130), 1 P[UCLA].

South Auckland: Coromandel, 26 Apr 1964, N (NZ 222), 15 L[UCLA]. Te

Puke, Prew Orchard, 31 Mar 1964, S (NZ 142), 19, 670 L, 23 P[UCLA].

Gisborne: Waihirere Domain, 25,26 Dec 1963, BS (NZ 47), 6lpo& (47-

102-107), 1 p? (47-101), 1¢, 16 L, 1 P[UCLA].

Nelson: Nelson City (Graham 1939:212).

TRIBE SABETHINI

Genus MAORIGOELDIA

1930. Maorigoeldia Edwards, 1930:302. TYPE SPECIES: *Culex argyropus

Walker, 1848, New Zealand; original designation (as subgenus of Rachi-

onotomyia).

Zo. Maorigoeldia argyropus (Walker)

Figs. 24, 41

1848. Culex argyropus Walker, 1848:2. TYPE: *Holotype 2, New Zealand;

identified as the type by Waterhouse [ BMNH ].

Maorigoeldia argyropus of Belkin (1962:492-494); Stone (1963:120); Pillai (1965).

Tripteroides (Maorigoeldia) argyropus of Edwards (1932:75); Taylor (1934:11);

Lee (1944:17); Stone, Knight and Starcke (1959:65).

Tripteroides (Maorigoeldia) argyropa of Lee (1946:229-230).

Tripteroides argyropus of Miller (1950:45); Miller and Phillipps (1952:27-28).

Rachionotomyia (Maorigoeldia) argyropus of Edwards (1930:302).

Rachionotomyia argyropus of Edwards (1924:360); Graham (1929:227-243; 1939:

213).

Uranotaenia(?) a argyropus of Theobald (1901:264-266).

Belkin: Culicidae of New Zealand 107

FEMALE (fig. 24; Pillai 1965:fig.1; Miller and Phillipps 1952:figs. 48-50;

Lee 1946:plate XIIla; Graham 1929:figs. 50-56). Wing: 4.25 mm. Proboscis:

2.84 mm. Forefemur: 2.67 mm. Abdomen: about 2.83 mm. As figured here

and as described by Belkin (1962:492, 494); with the following diagnostic fea-

tures. A beautiful large dark mosquito with striking silvery ornamentation with

very light bluish tinge as follows: a narrow orbital line on head; a broad dorsal

band involving apex of segment 2 and base of segment 3 of the palpus; a broad

lateral mesonotal stripe from humeral angle to wing root, interrupted at scutal

angle; a narrow continuous line from lower apn diagonallyto lower mep; patches

on ppl and upper part of pcx, on lower caudal part of stp, and in front of upper

mep bristles, not in upper stp as shown by Pillai (loc. cit.); patches on coxae

and trochanters; more or less complete postmedian and preapical rings on fe-

mora; anterior apical patch on tibiae, largest on hindleg; dorsal patch on apex

of segment 3, entire dorsal and anterior surfaces of segment 4 and anterior

base of segment 5 on foretarsus; on at least anterior surface from apex of seg-

ment 2 distad on midtarsus; from apex of segment 3 distad on hindtarsus; api-

colateral patches on abdominal tergites II-VII. Proboscis slightly swollen and

recurved in distal third in dry specimens. Palpal segmentation indistinct and

variable, segment 1 short and broad, segment 2 long and as broad as 1, seg-

ment 3 ankylozed with 2 and narrower and usually longer, a minute apical knob-

like segment 4 sometimes present. Spiracular bristles usually 2-4; always at

least 1 upper stp bristle. Acrostichals, dorsocentrals, prescutellars and su-

praalars numerous and strong. All legs with 2 subequal claws; all claws sim-

ple and with fine spicules on outer surface.

FEMALE GENITALIA (fig. 24). As figured. Genital segments deeply re-

tracted. Tergite VIII about 0.5 of VII, partially telescoped into latter, sparsely

covered with strong bristles distally. Sternite VIII prominent, subequal to

tergite, densely covered with heavy bristles except basolaterally where scales

are present. No intersegmental sclerotizations between segments VIII and IX.

Insula strongly developed and heavily sclerotized, long midventrally, with a few

short slender bristles laterally, continuous with sigma which is not joined to

cowl. Tergite IX about 0.35 of tergite VIII, moderately sclerotized, with about

3 Short heavy bristles near caudal margin on each side of midline. Cowl strong-

ly sclerotized, narrowed laterally, without setae; atrial plate strongly sclero-

tized ventrally, practically meeting its mate on midline, very lightly sclero- |

tized at its junction with cowl. Tergite X distinctly sclerotized, about 0.4 of

tergite IX, without setae. Postgenital plate separated by membrane from cowl,

with 2 widely separated lateral bristly lobes. Cercus short, about as broad as

wide, with many short bristles except on mesal surface. Spermathecae 3, one

very Slightly smaller than other 2.

MALE (fig.24; Graham 1929:fig.57). As figured here and as described

by Belkin (1962:492, 493), essentially as in the female except for sexual dif-

ferences. Palpal segment 4 present or absent as in the female; antennal whorls

much longer and denser than in female, flagellar segments 12 and 13 elongate.

Both claws of hindleg present and small; claws of foreleg and midleg enlarged,

unequal, simple. Proboscis more distinctly swollen and recurved apically than

in female.

MALE GENITALIA (Belkin 1962:fig.361; Lee 1946:fig.1; Graham 1929:

fig.58). As figured and described by Belkin (1962:492, 493). Segment IX very

strongly developed; tergite with prominent very broad median caudal lobe with

a minute median emargination and with very numerous Setae in several rows;

sternite long and broad. Sidepiece with specialized tergal bristles; basal mes-

al lobe strongly developed. Clasper simple, curved. Aedeagus complex; broad

108 Contrib..Amer. ‘Ent. Inst... vol. 3,°no. 1, 1968

at base; opisthophallus strongly developed, appearing as a broad lobe dorsad

of aedeagus proper (see discussion of male genitalia under Dixinae). Proctiger

very broad, paraproct with 2,3 small apical denticles on the main large tooth;

cercal sclerite strongly developed, cercal setae 6-8.

PUPA (Belkin 1962:fig. 361; Graham 1929:figs. 73-75). Abdomen: 3.25 mm.

Trumpet: 0.7mm. Paddle: 0.88 mm. As figured and described by Belkin

(1962:492, 493-494) with the following diagnostic features. Trumpet without

tracheoid. Paddle produced on apex, without hairs 1,2-P. Hairs 9-VIU, VIII

very large, subequal. The following additions to the chaetotaxy figured by Bel-

kin (1962) should be made on the basis of more adequate material. Hair 3-I

single, subequal to 12-C; 6-I single, subequal to 3-I; 5-II single, subequal to

7-I but thinner; 1-II usually 4, 5b (3-6); 3-III single, subequal to 3-II; 4-III usu-

ally single, subequal to 4-II; 5-III usually single or double (1-4), subequal to

5-II; 1-IV double or triple, subequal to I-III; 4-IV usually single, subequal to

4-II; 5-IV usually double, subequal to 5-V; 1-V usually single, about 0.75 of

tergite VI; 1-VI single, subequal to 3-VII; 3-VI single, subequal to 3-V; 5-VI

single, reaching to basal 0.2 of tergite VIII; 1-VII single, a little longer and

heavier than 4-VII; 9-VII subequal to 9-VIII but usually with about 10 branches

(7-13) instead of 13 (11-17).

LARVA (Belkin 1962:fig. 362; Miller and Phillipps 1953:fig.51; Lee 1944:

plate 4; Graham 1929:figs.62-72). Head: 1.35 mm. Siphon: 1.2 mm. Anal

Saddle: 0.2 mm. As figured and described by Belkin (1962:492, 494), with the

following diagnostic features and corrections. A very large larva with light

brown head capsule; thorax and abdominal segments 1-IV(V) whitish, (V)VI-VIII

yellowish; siphon and anal saddle darker than head capsule; gills sausage-

shaped, rounded apically and usually longer than siphon (not as figured by Bel-

kin). Head of very unusual shape (not as figured by Belkin from flattened speci-

men), distinctly wider than long (1.2), truncate in front and nearly semicircular

in outline in back, foramen magnum on ventral surface when viewed flat in a

mount. Comb scales in a very large patch. Siphon with distinct pecten and

regular paired rows of ventral and subdorsal hairs, mostly double. Anal saddle

widely incomplete; saddle hair (1-X) single, about twice length of siphon; ven-

tral brush composed of a single pair of long double hairs.

SYSTEMATICS. M. argyropus is one of the most clearly marked sabethine

species in the world. In the full development of the mesonotal bristles in the

adults, it has retained the primitive ancestral condition not found in any other

Old World sabethine. As I have indicated elsewhere (Belkin 1962:487-488), it

is readily separated from all other groups of Old World sabethines in all stages

and should be recognized as forming a distinct monotypic genus although it has

a number of similarities with members of the genus Tripteroides.

I can find no close relative of argyropus. Tripteroides (Rachionotomyia)

tasmaniensis (Strickland, 1911) from Southeast Australia and Tasmania has a

pattern of ornamentationof the thorax, legs and abdomen suggestive of argyropus

but it differs from the latter in important features of the male and larva. How-

ever, I think that argyropus probably represents the earliest offshoot of the

same lineage which gave rise to the monotypic tasmaniensis group and the

South Pacific caledonicus group of the subgenus Rachionotomyia of Tripteroides.

The most obvious variation I have noted in argyropus is in the segmenta-

tion of the palpus in both males and females. There is no indication of geo-

graphical differentiation in argyropus.

BIONOMICS. The natural breeding sites of argyropus were unknown until

D. A. Schroeder collected this species during our survey in February and March

Belkin: Culicidae of New Zealand 109

1964 in rot-holes in Nothofagus trees at Lake Rotoiti, Nelson (NZ 119) and on

Little Barrier Island (NZ 133), and Pillai (1965) discovered them ina knot-hole

in dead Nothofagus fusca in South Otago in May 1965. Natural breeding sites

seem to be uncommon and difficult to locate for collectors but fortunately this

interesting species has adapted readily to breeding in just about any kind of ar-

tificial receptacle containing water in the vicinity of native forests, including

water tanks. One of our collections was made ina drying stream bed pool with

leaves on Little Barrier Island (NZ 128). In this collection and 3 others in tins

and a tire (NZ 61, 63, 191B), argyropus was associated with Aedes notoscriptus.

Graham (1929:227-243) gives extensive notes on this species, including the

description of the egg and oviposition. The egg is said to be ovoid, laid singly

on the water surface and to split lengthwise near the ventral surface in eclo-

sion. The larvae have the usual sabethine habit of resting on the bottom of the

breeding container with the dorsal surface down. They seldom come to the

surface film to obtain air and are very hardy.

All the adults I have seen were collected during the months of December

through February. However, young larvae were obtained in July at Titirangi

Beach (NZ 191B) ina small tin together with Aedes notoscriptus, females of

which were biting at the time. Graham (1929:232) reports "hibernating" argy-

ropus adults under dead nikau and tree fern fronds and in week-end cottages in

the Waitakere Ranges. I think, therefore, that at the latitude of Auckland at

least, both adults and immature stages survive through the winter and breeding

may continue at a slower rate. Pillai (1965:34) implies that in South Otago

adults die in the fall after laying eggs towards the end of the autumn and a new —

crop appears during the spring from the overwintering larvae.

Graham (1929:232) states that argyropus females are persistent biters.

This was not our experience in areas of fairly high larval density where we

saw a number of adults resting on tree trunks and warily approaching in the

typical flying attitude of sabethines but never landing or biting. The relative

rarity of the strikingly marked adults of this species in collections also supports

the view that argyropus is not readily attracted to man. Pillai (1965:35) notes

the absence of this species in mosquito pools for arbovirus studies in areas

where the immature stages have been collected.

DISTRIBUTION (fig.41). Material examined: 409 specimens; 33 0, 2592,

325 larvae, 26 pupae; 14 individual rearings (10 larval, 1 pupal, 3 incomplete).

NEW ZEALAND. North Auckland: Waitakeres, 18 Feb 1959, M. Knight,

19[AUCK]. Titirangi, 2 Feb 1945, G.L. Bissett (588), 1 L, 2 P[UCLA], 2 L,

1 P [NELS]; locality questionable (557), 4 L[NELS], (605), 1%, 19 [NELS].

Titirangi Beach, end of Mahoe Road, 30 Dec 1964, BF (NZ 61B), 3 lpo& (61B-

102,104,107), 4 1p? (61B-105, 106, 117,118), 2 1p (61B-103, 121), 50 L [UCLA];

same data (NZ 63), 40°, 49, 17 L[UCLA]; same locality, 11 Jan 1964, BS

(NZ 90), 2 [UCLA]; same locality, 4 July 1964, F (NZ 191B), 16 L [UCLA].

Clevedon, 5 Apr 1964, S(NZ 143), 11 L; same data (NZ 144), 1-po (144-101),

GC. 6 8 04h TP POCLAT:

Little Barrier Island: Small stream between Te Waikohare and Tirikikawa

streams, 6 Mar 1964, S (NZ 128), 24 L[UCLA]. Summit Track, elev. 750 ft,

7 Mar 1964, S (NZ 133), 2 1lpo (133-101, 104), 1 1p? (133-103), 1 1p (133-102),

18 L [UCLA].

Wellington: Ohakune, elev. 2060 ft, Feb 1920, T.R. Harris, 1 o [BMNH];

same elev., 24 Jan 1922, T.R. Harris, 1 9 [USNM]; elev. not specified, 15

Dec 1922-15 Jan 1923, T.R. Harris, 3 o [BMNH]; Jan 1924, T.R. Harris, 1 ¢

[BMNH], 1“ [USNM]; Feb 1924, T.R. Harris, 1 oc [BMNH]. Wellington, Wil-

110 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

ton Bush, Feb 1922, on tree trunks, G.V. Hudson, 1, 2 2 [BMNH]; locality

and date not specified, Hutton (118C), 1 o [CANT].

Nelson: Nelson, north branch Maitai River, elev. ca 400 ft, 11 Feb 1964,

S (NZ 123A), 1 o [UCLA]. Aniseed Valley, 1-4 Dec 1923, A.L. Tonnoir, 4¢%

[NELS], 1 o [CANT]. Lake Rotoiti, 9 Feb 1964, S(NZ 119), 29 L[UCLA].

Marlborough: Goose Bay, 4 Feb 1925, A.L. Tonnoir, 1% [CANT].

Canterbury: Price's Bush, Banks Peninsula, 2 Dec 1934, S. Lindsay, 1 ¢,

1 9 [CANT].

Westland: Whataroa, Scenic Reserve, July 1964, L (Pillai 1966). Okarito

Lagoon, June 1965, L (Pillai 1966).

Otago: Southeastern coast, lat. 46935'S, May-Sept 1965, L (Pillai 1966).

New Zealand, locality not specified: Holotype ? [BMNH]; 1 2? [WELL].

SUBFAMILY CHAOBORINAE

The Phantom Midges

The phantom midges and related forms are readily recognized from true

mosquitoes and dixa midges by the characters given in the keys to subfamily.

The general features of the Chaoborinae have been reviewed by Belkin (1962:

535-537). The phallosome in this subfamily is very simple, consisting of a

poorly developed basal piece and a pair of aedeagal sclerites articulating di-

rectly with it, without a distinct intermediate paramere. Apparently no aedea-

gal pouch is ever developed in this group and the aedeagus is therefore erect at

rest and not retracted.

The Chaoborinae are all predaceous in the larval stage. They are best

represented on large continental areas, especially in the Holarctic region, but

the tribes Corethrellini and Chaoboriniare known from relatively small islands.

To date, the very primitive Corethrella novaezealandiae, for which a new sub-

genus is here recognized, is the only representative of the subfamily reported

from New Zealand. It is surprising that no member of the tribe Chaoborini

has been found in New Zealand. It is quite possible that this tribe is also rep-

resented but has been overlooked in New Zealand because of the midgelike ap-

pearance of the adults of the smaller species and the specialized breeding

sites of the immature stages. A thorough survey for Chaoborus-like forms

should be made with light traps for adults and dredging nets for immature

stages in lakes and ponds.

TRIBE CORETHRELLINI

Genus CORETHRELLA

1902. Corethrella Coquillett, 1902:191. TYPE SPECIES: *Corethra brakeleyi

Coquillett, 1902, New Jersey, U.S.A.; original designation.

1911. Ramcia Annandale, 1911:187. TYPE SPECIES: *Ramcia inepta Annandale,

1911, Ceylon; monobasic.

1942. Lutzomiops Lane, 1942:127. TYPE SPECIES: Corethrella nigra Lane,

1939, Brazil; original designation.

Notocorethrella Belkin, n. subg.

Belkin: Culicidae of New Zealand 111

TYPE SPECIES: Corethrella novaezealandiae Tonnoir, 1927.

ADULTS. Vestiture of body and legs entirely of hairs and bristles, no dis-

tinct scales developed; bristles moderate in length. Pleural bristles on upper

and lower apn, upper ppn and upper mep. Claws all toothed in the male, prob-

ably also in female (not noted). Vein Ry ending on C well beyond furcation of

Ro.43, distance between apex of R2 and Rj only slightly more than that between

apex of Sc and R1; Rs only slightly longer than crossvein r-m; vein M beyond

crossvein m-cu only slightly shorter than vein Mj,9; base of Cu, forming an

angle of more than 60° with base of Cug. Vestiture of all wing veins except C

entirely of long hairs without any indication of striations; vein C and fringe

with flat broad striated scales in addition to hairs.

MALE GENITALIA. Sidepiece without specialized thickened bristles near

base but with several thickened short bristles on mesal and sternomesal sur-

face distally. Clasper distinctly shorter than sidepiece, broad, strongly ex-

panded in distal half; with a distinct short, heavy spiniform and several minute

setae near apex; long mesal seta submedian in position. Ninth tergite very

large.

24. Corethrella (Notocorethrella) novaezealandiae Tonnoir

Figs. 3, 42

. 1927. Corethrella novae-zealandiae Tonnoir, 1927:107-108. TYPE: *Holotype

?, Otira (Westland), New Zealand, 7 Feb 1922, A.L. Tonnoir [NELS].

Corethrella novaezealandiae of Edwards (1932:19); Taylor (1934:8); Miller

(1950:43); Belkin (1962:540).

FEMALE (Tonnoir 1927:fig.2; Belkin 1962:fig.404). Wing: 2.5mm. En-

tire body: 2.0 mm. As described by Tonnoir and Belkin, with the characters

of the subgenus and the following diagnostic features. General coloration dark

yellowish; body and legs with hairs only. Wing slightly yellowish and with in-

fuscations at the crossveins, furcations, in cell C and in apex of cell Sc; veins

with golden hairs; fringe metallic pale golden.

MALE (fig.3). As figured; essentially as in the female except for anten-

nal and leg characters; additional morphological details described from slide

mounts are probably applicable to both sexes; bristles of mesonotum probably

longer than figured from slide-mounted specimen. Antennal scape smaller than

in female, with 2, 3 bristles; torus enlarged; bristles of basal flagellar whorls

longer but not very numerous (about 8); proportions of flagellar segments as

figured. Palpus distinctly 5-segmented, all segments with setae; segments I

and 2 short, 2 a little longer, segment 3 about 2.5 of 2, segment 4 about 1.3 of

2, segment 5 a little longer than 3. Upper part of apn with about 5 short thick-

ened bristles, lower part with about 4 longer thinner bristles; upper mep with

1 long and 1 short bristle; metanotum with 3 bristles on each side. Legs ap-

parently with only a few extremely long hairs which are usually developed in

most species of Corethrella in greater number and larger size on the femora,

tibiae and first tarsal segments. Claws of foreleg and midleg greatly enlarged,

distinctly longer than tarsal segment 5, strongly recurved, members of a pair

not markedly different in size but mesothoracic longer than prothoracic; all

with a slender sharp ventral tooth in basal half but without outer spicules; an-

terior claw on both legs with a long slender spine projecting outward from near

112 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

base (probably homolog of outer spicules); hindclaws much smaller, subequal

in size, with a short sharp ventral submedian tooth, anterior claw with a short

curved spine projecting from outer surface; probably hindclaws similar to those

of female. Wing venation and vestiture as figured; probably scales on vein C

more numerous than shown but fringe appears to be much less developed than

usual in other species. Abdomen with minute spiracles on segments II-VII.

MALE GENITALIA (fig.3). As figured and with the diagnostic features

indicated for subgenus. Tergite IX very large, with numerous setae some of

which project beyond apex of sidepiece. Sidepiece with long bristles laterally

and ventrally toward apex; base of dorsal surface without conspicuous setae;

middle of tergomesal area with 6-8 slender setae, followed ventrodistad by 3

thickened longer straight setae; distal sternomesal margin with 2 or 3 similar

but stronger setae with more prominent basal tubercles. Clasper short, broad

and further expanded in distal two-thirds in dorsal plane; a large seta near the

inner sternal margin at about 0.45 from base; a distinct short heavy spiniform

near sternal end of broad apex, followed dorsad by a line of about 6 minute se-

tae. Aedeagus (penis valves) bulbous at base, slender and only slightly taper-

ing in distal half beyond the neck.

IMMATURE STAGES. Unknown.

SYSTEMATICS. C. novaezealandiae is the only representative of the sub-

family found to date in New Zealand. In common with other culicids of New

Zealand it appears to have retained several important primitive features of its

phyletic line, notably the following: (1) vestiture of the body, legs and wing

veins other than C entirely of bristles and hairs, without any indication of flat-

tening or striations, (2) ends of veins Sc and Ry on costa widely separated, (3)

fringe of wing relatively poorly developed, (4) sidepiece of male genitalia with-

out marked specializations, (5) clasper relatively short and with a well devel-

oped spiniform, and (6) tergite IX very strongly developed in the male. Since

this combination of features separates novaezealandiae very distinctly from all

other species of Corethrella known to me, I am proposing a new subgenus for

this unique form. Should the immature stages of novaezealandiae prove to be

as distinctive as its adults, Notocorethrella may have to be raised to full gen-

eric rank.

BIONOMICS. This species probably breeds in the grassy margins of lakes

and ponds as do many other species of Corethrella. To date only 2 males and

4 females have been collected, all in Westland. The 2 males were taken ina

landing-biting collection on the shore of Lake Mapourika just after dark.

DISTRIBUTION (fig.42). Material examined: 2c, 49. NEW ZEALAND.

Westland: Lake Brunner, 2 Feb 1922, A.L. Tonnoir, 12[NELS]. Otira, 7 Feb

1922, A.L. Tonnoir, 2 holotype [NELS]. Lake Mapourika, 6 Feb 1964, S (NZ

112), 1 o (112-1) [NELS],.1.o (112-2) [UCLA]... Waiho, 30 Jan 1922, A. L.

Tonnoir, 1¢[NELS], 1 2 [BMNH].

Belkin: Culicidae of New Zealand 113

RECORDS OF EXTRALIMITAL SPECIES

Graham (1939) reported the following extralimital nominal species from

North Auckland. I have not Seen any of the material on which these records are

based and no additional records of any of these forms have been reported.

Anopheles maculipennis Meigen, 1818 was identified by Graham (1939:213)

on the basis of 1 live female taken at Auckland on a ship from the East Indies

on 27 May 1929 and another female taken on 4 Sept 1929 on a ship from Samar-

ang. This is obviously a misidentification as the maculipennis complex is not

represented in the East Indies. There is no way of determining the actual spe-

cies seen by Graham.

Culex annulirostris Skuse, 1889 was discovered by Graham (1939:111)

breeding in the hold of S.S. Tofua on arrival at Auckland from Suva and subse-

quently breeding ina barrel on the waterfront. This is a plausible identifica-

tion as this species may be dispersed in boats and possibly ships in the tropical

South Pacific. However, Graham's identifications of immature stages are not

reliable and it is possible that the larvae were actually pervigilans.

Aedes vexans (Meigen, 1830) was identified by Graham (1939:213) on the

basis of a single collection of larvae found "in a tin of water jammed among

rocks just above high tide at Russell in July, 1929."' I think it is very likely

that the species involved was Aedes (F.) notoscriptus which is known from this

general area, utilizes containers for breeding and tolerates a fair degree of

salinity (collections NZ 37, 38).

114 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

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Forsyth, D.

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1965. Colonization and biology of wa fuscus. Roy. Soc. New Zeal.,

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1956. Report on the Diptera of Auckland and Campbell islands. Dominion

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116 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

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1954. A mosquito survey in New Caledonia and the Belep islands, with new

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120 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

FIGURES

Paradixa neozelandica, female

Opifex fuscus, male

Corethrella novaezealandiae, male

Nothodixa campbelli, male and female

Nothodixa campbelli, larva and pupa

Nothodixa septentrionalis, larva and pupa

Nothodixa otagensis, Nothodixa septentrionalis, Paradixa fuscinervis,

Paradixa tonnoiri, Paradixa harrisi, male genitalia

8. Paradixa neozelandica, male and female

9. Paradixa neozelandica, larva and pupa

10. Paradixa fuscinervis, larva and pupa

11. Paradixa tonnoiri, larva and pupa

12. Paradixa harrisi, larva and pupa

13. Culex rotoruae, male genitalia and pupa

14. Culex asteliae, male genitalia and pupa; Culex pervigilans, male genitalia

15. Culex asteliae, larva

16. Coquillettidia Coquillettidia tenuipalpis, male and female

17. Coquillettidia tenuipalpis and Coquillettidia iracunda, pupae

18. Opifex fuscus, male and female

19. Aedes chathamicus, male and female

20. Aedes chathamicus, male genitalia and pupa

21. Aedes chathamicus, larva

22. Aedes subalbirostris, male genitalia and pupa

23. Aedes subalbirostris, larva

24. Maorigoeldia argyro} argyropus, male and female

25. Distribution of Nothodixa campbelli and Nothodixa philpotti

26. Distribution of Neodixa minuta, Nothodixa otagensis and Nothodixa septen-

trionalis

27. Distribution of Paradixa neozelandica

28. Distribution of Paradixa fuscinervis

29. Distribution of Paradixa harrisi and Paradixa tonnoiri

30. Distribution of Culex pervigilans, general

31. Distribution of Culex pervigilans, specific

32. Distribution of Culex asteliae and Culex rotoruae

33. Distribution of Culex quinquefasciatus

34, Distribution of Culiseta tonnoiri

35. Distribution of Coquillettidia tenuipalpis

36. Distribution of Coquillettidia iracunda

37. Distribution of Opifex fuscus, Aedes chathamicus and Aedes australis,

IOonh WH ke

general

38. Distribution of Opifex fuscus, Aedes chathamicus and Aedes australis,

specific

39. Distribution of Aedes antipodeus and Aedes subalbirostris

40. Distribution of Aedes notoscriptus

41. Distribution of Maorigoeldia argyropus

42. Distribution of Corethrella novaezealandiae

fy a ae ee

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LAMBERT CONFORMAL CONIC PROJECTION

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400 600

154 156 58 . 160 162 164 166 168 170 172

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SOUTH PACIFIC- Sheet 5

LAMRERT CONFORMAL CONIC PROJECTION

STANDARD PARALLELS 33° and 45°

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STATUTE MILES

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CORETHRELLA

e novaezealandiae

Belkin: Culicidae of New Zealand 163

APPENDIX A

NZ Collection Data

The bulk of the material on which this study is based was collected and

reared following the methods later outlined by Belkin, Hogue et al (1965). This

material is identified only by the following printed label: NZ [handwritten col-

lection number from 1-225] 63/New Zealand 64/Belkin/Schroeder/. All the

data pertinent to each collection (locality, date, collector, breeding site,

rearing information, etc.) are entered on record cards deposited in the Depart-

ment of Zoology, University of California, Los Angeles. These data are sum-

marized in the following list under each collection number. Included are a few

miscellaneous collections bearing this general label, which may not contain

culicid material or were made by other individuals subsequent to the main sur-

vey. The following abbreviations are used for the names of collectors: B(J.N.

Belkin); D (L.J. Dumbleton); E (R. Elliott); F (D.A. Forsyth); G (E.S. Gour-

lay); N(E.R. Nye); S(D.A. and M.R. Schroeder).

1. Epsom, Auckland, Golf Road (North Auckland), 21 Aug, S. On wall in

house; 2000 hrs. Tipulid.

2. Waitomo Caves, 4 mi E (South Auckland), 24 Aug 63, S. On outside of

car window; 1200 hrs. Chironomid.

3A. Kaiapoi, Pineacres Motel (Canterbury), 27 Aug 63, S. Inside bath-

room; 1930 hrs. Chironomids.

3B. Kaiapoi, Pineacres Motel (Canterbury), 28 Aug 63, S. Inside bath-

room; 0700 hrs. Culex (C.) pervigilans; chironomids.

4, Westport, Luxury Motel (Nelson), 28 Aug 63, S. On wall of bathroom;

2130 hrs. Chironomids.

5. Mt. Albert, Auckland, DSIR Laboratory (North Auckland), 10 Sept 63,

A. Watson. In laboratory; 1100 hrs. Tipulid.

6. Rangitoto Island, Summit Track (North Auckland), eley. 177 m, 14 Sept

63, S. Cement container, 1 ft diameter, 3 ft deep; decaying vegetation; par-

tial shade. Aedes (F.) notoscriptus.

7. Rangitoto Island (North Auckland), 14 Sept 63, S. Leaf axils of aste-

lias (Collospermum hastatum) on ground in native bush; reddish brown water,

partial shade. Culex (C.) asteliae; chironomid.

8. Rangitoto Island (North Auckland), 14 Sept 63, S. Old cement founda-

tion for gun emplacement in native bush; fresh water with algae and decaying

vegetation. Aedes (F.) notoscriptus.

9. Northcote, Clarence Road (North Auckland), elev. 12 m, 30 Sept 63, S.

Metal tank, 5 ft diameter, 2.5 ft deep; greenish brown water with scum, algae

and water lilies; partial shade. Culex (C.) pervigilans; chironomid.

10. Little Barrier Island, Ranger's house (Hauraki Gulf), elev. 6 m, 9 Oct

63, S. Metal pan, 2 ft diameter, 1 ft deep, 1.5 m above ground; clear water

with leaves; partial shade. Culex (C.) pervigilans.

11. Little Barrier Island, small stream between Te Waikohare and Tiriki-

kawa streams (Hauraki Gulf), elev. 7m, 9 Oct 63, S. Stream bed pool with

fresh water and no vegetation; partial shade in native bush. Paradixa fusci-

nervis; Culex (C.) pervigilans; Aedes (O.) antipodeus.

164 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

12. Little Barrier Island, Summit Track (Hauraki Gulf), elev. 267 m, 9

~ Oct 63, S. Ground pool, 12 x 3 ft; brownish fresh water with scanty grass on

margins and decaying leaves; partial shade in native bush. Culex (C.) pervigi-

-lans; Aedes (O. ) antipodeus.

13. Little Barrier Island, Summit Track (Hauraki Gulf), elev. 186 m, 10

Oct 63, S. Leaf axils of astelias growing on ground in native bush; water red-

dish brown, with decaying leaves; partial to deep shade. Culex (C.) asteliae.

14. Cuvier Island, Monument side (Hauraki Gulf), elev. 2 m, 24 Oct 63, S.

Seaside seepage on rocky shore; water clear, fresh and with decaying vegeta-

tion; partial shade. Opifex fuscus.

15A, B. Great Barrier Island, Shoal Bay (Hauraki Gulf), on board M. V.

Colville, 24-25 Oct 63, Captain Brown. Inside main cabin, at light (A), early

morning (B). Tipulids.

16. Great Barrier Island, Shoal Bay (Hauraki Gulf), elev. 2m, 25 Oct 63,

S. Ditch along dirt road; clear fresh water with abundant scum, algae and

grassy, floating and submerged vegetation; partial shade. Majority of larvae

developed whitish growth around siphon and died. Culex (C.) pervigilans.

17. Great Barrier Island, mouth of Te Wairere stream (Hauraki Gulf),

elev. 9m, 26 Oct 63, S. Seaside rockhole with clear, slightly brackish water

with some scum, algae and a few decaying leaves; partial shade. Culex (C. )

pervigilans; Opifex fuscus.

18A, B,C. Little Barrier Island (Hauraki Gulf), elev. 7m, 29 Oct (A), 7

Nov (B), 9 Nov (C) 63, S. Inside bunkhouse; 2000 hrs (A), daylight (B), 0630

hrs (C). Aedes (O. ) antipodeus (ALC): svoetbphilid (B).

19A, B. Little Barrier Island (Hauraki Gulf), elev. 7m, 30 Oct (A), 1 Nov

(B) 63, S. In lavatory of Ranger's house, 2230 hrs. Aedes (O.) antipodeus.

20. Little Barrier Island, between Lamb Bay Creek and Awaroa Stream

(Hauraki Gulf), elev. 5 m, 31 Oct 63, S. Rocky stream bed pool near shoreline

cliffs; clear fresh water with algae and decaying leaves; partial shade in native

bush. Culex (C.) pervigilans.

21. Little Barrier Island, 0.25 mi up Awaroa stream (Hauraki Gulf), elev.

ca30m, 31 Oct 63, S. Adults around boulders and along bank of stream bot-

tom (no flowing water); 1430 hrs, partly cloudy.

22A. Little Barrier Island, about 0.25 mi up Awaroa Stream (Hauraki Gulf),

elev. ca30 m, 31 Oct 63, S. Adults found drowned on water surface of stream

bed pool (same as 22B). Culex (C.) pervigilans.

22B. Little Barrier Island, about 0.25 mi up Awaroa Stream (Herat Gulf),

elev. ca30 m, 31 Oct 63, S. Rocky stream bed pool; water fresh and with al-

gae; partial shade. Culex (C.) pervigilans.

23A. Little Barrier Island, 0.25 mi up Awaroa Stream, at junction of small

tributary stream (Hauraki Gulf), elev. 30 m, 31 Oct 63, S. Adults swarming

along bank behind stream bed pool of collection 23B; 1500 hrs; partly cloudy.

Culex (C.) pervigilans.

23B. Little Barrier Island, 0.25 mi up Awaroa Stream, at junction of small

tributary stream (Hauraki Gulf), elev. 30 m, 31 Oct 63, S. Stream bed pool

with clear fresh water and decaying leaves; paletial shade in native bush. Culex

(C.) pervigilans.

24. Little Barrier Island, Summit Track (Hauraki Gulf), elev. 267m, 8

Nov 63, S. Leaf axils of astelias growing on ground; water with decaying

leaves; partial to deep shade in native bush. Culex (C.) asteliae.

25. Little Barrier Island, Summit Track (Hauraki Gulf), elev. 186m, 8

Nov 63, S. Same site as collection 13. Leaf axils of epiphytic astelias (3.5 m

Belkin: Culicidae of New Zealand 165

above ground); water reddish brown and with decaying leaves; partial shade in

native bush. Culex (C.) asteliae.

26. Little Barrier Island, Summit Track (Hauraki Gulf), elev. 75 m, 8 Nov

63, S. Leaf axils of astelias growing on ground. Culex (C.) asteliae.

27. Cape Kidnappers (Hawke's Bay), elev. 66m, 16 Nov 63, S. Stream

bed pool with silty, turbid fresh water with slow current, abundant vegetation

and mud bottom; full sunlight; sheep pasture. Culex (C.) pervigilans.

28. Ormond, Waimare Orchard (Gisborne), elev. 25 m, 17 Nov 63, S. Ce-

mented fish pond with clear fresh water and abundant cultivated plants; partial

shade. Culex (C.) pervigilans.

29. Near Okiore, 0.5 mi downstream from midway point in Waioeka Gorge

(Gisborne), 19 Nov 63, S. Stream margin rockhole with clear fresh water,

abundant algae and rocky and sandy bottom; full sunlight. Culex (C.) pervig-

ilans.

30. Epsom, Auckland, Golf Road (North Auckland), 23 Nov 63, S. In

rearing room, probably escaped from rearing vial.

31. Epsom, Auckland, Golf Road (North Auckland), 28 Nov 63, S. Landing

on front porch, at light; 2300 hrs; cloudy night. |

32. Epsom, Auckland, Golf Road (North Auckland), 1 Dec 63, S. At light

on front porch, 2230 hrs; clear moonlit night; slight wind.

33. Little Barrier Island (Hauraki Gulf), 7 Nov 63, S. Collection record

uncertain, probably as stated. Aedes (O.) antipodeus.

34. Little Barrier Island (Hauraki Gulf), elev. 7m, 10 Oct 63, S. Inside

lavatory of Ranger's house, at light, 2100-2300 hrs.

390. Waitakere Ranges, Quarry tributary of Waitakere River, at edge of

Cascade Park, bridge at intersection (North Auckland), 18 Dec 63, BS. Stream

margin; slow current; clear fresh water with abundant flotage and grassy veg-

etation; mud bottom; full sunlight. Paradixa neozelandica; P. fuscinervis;

Culex (C.) pervigilans.

36. Ruakaka, 1 mi S on highway 1 (North Auckland), 20 Dec 63, BS. Pond

with clear fresh water; grassy vegetation abundant; mud bottom; full sunlight.

Paradixa fuscinervis; Culex (C.) pervigilans; Coquillettidia (C.) iracunda.

37. Opua, 3 mi S of turnoff on Kawakawa-Paihia road (North Auckland), 20

Dec 63, BS. Small treeholes in mangrove (Avicennia) with dark reddish brown

water; holes from a few inches to 1 ft above high tide level; water apparently

fresh. Aedes (F.) notoscriptus.

38. Opua, 3 mi S of turnoff on Kawakawa-Paihia road (North Auckland),

20 Dec 63, BS. Small treeholes in mangrove (Avicennia) with dark reddish

brown water, apparently fresh; holes 2-2.5 ft above high tide level. Culex (C.)

pervigilans; Aedes (F.) notoscriptus.

39. Kaeo, 3 miS on highway 10 (North Auckland), 20 Dec 63, BS. Small

pool in ditch along highway; fresh, turbid, milky water without vegetation; mud

bottom; partial shade. Culex (C.) pervigilans.

40. Mangamuka Gorge, 4 mi N of Soda Spring, NW drainage near top of

pass (North Auckland), 21 Dec 63, BS. Leaf axils of broad-leaved astelias (1-

3m above ground); water brownish; mud in axils; partial shade. No mosqui-

toes; chironomids present.

41. Mangamuka Gorge, 2 mi N of Soda Spring (North Auckland), 21 Dec 63,

BS. Stream bed pool along moderately fast-flowing stream; clear fresh water

without vegetation; clay bottom; partial shade. Nothodixa campbelli; Paradixa'

neozelandica; P. fuscinervis; Culex (C.) pervigilans.

42. Mangamuka Gorge, 2 mi N of Soda Spring (North Auckland), 21 Dec 63,

166 Contrib... Amer. .Ent.: Inst,, -vol.,3, no..4, 1968

BS. Margins of moderately fast-flowing stream, larvae on banks; clear fresh

water without vegetation; clay bottom; partial shade. Paradixa neozelandica;

P. fuscinervis.

43. Waipoua Forest, 1-2 mi N of Big Tree (North Auckland), 21 Dec 63,

BS. Leaf axils of epiphytic astelias (1-2 m above ground); water brownish;

axils with mud and decaying leaves and twigs; partial shade. Chironomids;

mosquitoes absent.

44. Waipoua Forest, Big Tree Roadside Park (North Auckland), 21 Dec

63, BS. Three separate holes in large tree stump near rest rooms, about 1.5

m above ground; water reddish brown; partial shade. One small mosquito lar-

va seen but lost.

45. Oratia, DSIR Field Station (North Auckland), 22 Dec 63, BS. Fire drum

with 6 in of rusty water; full sunlight. Paradixa fuscinervis; Culex (C.) per-

vigilans. woe

46. Rotorua, Kuirau Hot Springs Reserve (South Auckland), 23 and 27 Dec

63, BS. Pools and outflow ditch from hot springs; temperature 27-289C; water

turbid and with moderate current; vegetation scanty to none, herbaceous; mud

bottom; full sunlight. Paradixa fuscinervis (probably contamination); Culex

(C.) rotoruae.

47. Ormond, Waihirere Domain (Gisborne), 25 and 26 Dec 63, BS. Small

treehole in Leptospermum ericoides tree; dark reddish brown water; bottom

with decaying vegetation; partial shade. Aedes (F.) notoscriptus.

48. Ormond, Waihirere Domain (Gisborne), 25 and 26 Dec 63, BS. Stream

margin and pools above Swimming pool to falls; current moderate; water clear

and fresh; flotage abundant, vegetation scanty; mud bottom; partial shade.

Larvae frequently on leaves floating in stream pools or on banks of stream.

Paradixa neozelandica; P. fuscinervis.

49. Ormond, Waihirere Domain (Gisborne), 26 Dec 63, BS. Axils of epi-

phytic astelias (3-4m above ground) on upper part of path to falls; water brown-

ish; partial to deep shade in native bush. No mosquitoes.

50. Hexton, at bridge H20,S16-41 (Gisborne), 26 Dec 63, BS. Grassy

stream margins; water clear to turbid, fresh; current moderate; vegetation

scanty; mud bottom; full sunlight. Paradixa neozelandica; P. fuscinervis; Cu-

lex (C.) pervigilans. :

51. Terapatiki (Gisborne), 27 Dec 63, BS. Margins of small stream; cur-

rent moderate to strong; water fresh and clear; vegetation scanty; rock bot-

tom; full sunlight. Paradixa fuscinervis. :

52. Urewera National Park, Aniwaniwa Falls (Gisborne), 27 Dec 63, BS.

Stream pool below upper falls; current moderate; water fresh and clear; vege-

tation scanty, flotage abundant; gravel and rock bottom; partial to full sunlight.

Paradixa fuscinervis.

53. Urewera National Park, Aniwaniwa Falls (Gisborne), 27 Dec 63, BS.

Leaf axils of Cordyline australis (at 2.5 to 3 m above ground) on path to falls;

water turbid; partial shade. Ceratopagonids abundant.

54. Urewera National Park, Aniwaniwa Falls (Gisborne), 27 Dec 63, BS.

Leaf axils of epiphytic astelias (at 2-3 m above ground) on path to falls; water

brownish; partial to deep shade. Chironomids abundant.

55. Urewera National Park, near Mokau Falls (Gisborne), 27 Dec 63, BS.

Blocked road ditch, probably fed by seepages, along main road, current slow;

water brownish, fresh; grassy vegetation and flotage scanty; mud bottom; full

sunlight. Paradixa fuscinervis; Culex (C.) pervigilans.

56. Urewera National Park, Hopuruahine area (Gisborne), elev. ca 2900 ft,

Belkin: Culicidae of New Zealand 167

27 Dec 63, BS. Margins of small stream; current strong; water clear and

fresh; scanty grassy vegetation; gravel and rock bottom; deep shade. Notho-

dixa septentrionalis.

o1A. Waitakere Ranges, Bethells Road Swamp, near mouth of Waitakere

River (North Auckland), 28 and 29 Dec 63, BFS. Edge of swamp. Biting-land-

ing and Sweeping collection; 1130-1445 hrs; sunny, warm and windy. Coquillet-

tidia (A. ) tenuipalpis; C.(C.) iracunda.

07B. Waitakere Ranges, Bethells Road Swamp, near mouth of Waitakere

River (North Auckland), 28 and 29 Dec 63, BFS. Shallow water along edge of

Swamp; water stagnant, fresh, brownish; vegetation very dense, primarily Ty-

pha, Juncus, Scirpus and sedges; bottom with mud and decaying vegetation; full

sunlight. Coquillettidia (A.) tenuipalpis; C.(C.) iracunda.

08. Waitakere Ranges, Quarry tributary of Waitakere River, at edge of

Cascade Park, downstream of bridge at intersection (North Auckland), same

location as collection 35 but farther downstream, 28 Dec 63, BF. Stream mar-

gin; Slow current; clear fresh water with abundant flotage and grassy vegeta-

tion; mud bottom; deep shade. Paradixa fuscinervis.

09. Waitakere Ranges, Quarry tributary of Waitakere River, at edge of

Cascade Park, downstream of bridge at intersection (North Auckland), same

location at collection 35 but farther downstream, 28 Dec 63, BF. Biting-land-

ing in dense shade along stream banks. Coquillettidia (C.) iracunda; Aedes (F.)

notoscriptus.

60. Waitakere Ranges, Quarry tributary of Waitakere River, at edge of

Cascade Park, downstream of bridge at intersection (North Auckland), same

location as collection 35 but farther downstream, 28 Dec 63, BF. Leaf axils

of terrestrial astelias; water turbid, brownish; axils with mud; deep shade.

Aedes (F.) notoscriptus; chironomids.

61A. Titirangi Beach, end of Mahoe Road (North Auckland), 30 Dec 63,

BF. Adults biting-landing on collectors near breeding sites (61B); 1030-1130

hrs; intermittent rain. Aedes (F.) notoscriptus.

61B. Titirangi Beach, end of Mahoe Road (North Auckland), same location

and time as 61A, 30 Dec 63, BF. Tin cans and tire; water brownish; bottom

with decaying vegetation; deep shade. Aedes (F.) notoscriptus; Maorigoeldia

argyropus.

62. Titirangi Beach, end of Mahoe Road (North Auckland), 30 Dec 63, BF.

Leaf axils of epiphytic astelias (3-4 m above ground); water brown, turbid;

bottom with mud; deep shade. Mosquitoes absent.

63. Titirangi Beach, end of Mahoe Road (North Auckland), 30 Dec 63, BF.

Tin cans; water rusty, brown; deep shade. Aedes (F.) notoscriptus; Maori-

goeldia argyropus.

64. Titirangi Beach, creek above dam (North Auckland), 30 Dec 63, BF.

Margins of dammed creek; water fresh, clear; vegetation herbaceous, scanty;

flotage abundant; mud bottom; partial to deep shade. Paradixa neozelandica;

P. fuscinervis; Culex (C.) pervigilans.

65. Little Huia, west of point beyond pier (North Auckland), 30 Dec 63,

BF. Cavity in dead horizontal trunk of pohutukawa tree projecting over high

tide level; water turbid, brackish; some flotage and decaying vegetation; full

sunlight. Opifex fuscus.

66. Oratia, DSIR Field Station (North Auckland), same site as collection 45,

30 Dec 63, S. Fire drum with 6 in of rusty water; full sunlight. Paradixa fus-

cinervis; Culex (C.) pervigilans; C.(C.) quinquefasciatus.

67. Rangitoto Island (North Auckland), 4 Jan 64, S. Leaf axils of astelias

168 Contrib: Amer, Ent. Inst.) voli 3, no,-1, 1968

(Collospermum hastatum) growing on lava flow near the sea; water reddish

brown, fresh; axils with decaying leaves and twigs; partial shade. Culex (C.)

asteliae; Aedes (F.) notoscriptus.

| 68. Rangitoto Island (North Auckland), 7 Jan 64, BS. Leaf axils of astelias

(Collospermum hastatum) growing on periphery of clumps of vegetation on lava

flow (farther from shore than collection 67); water light reddish brown, fresh;

axils with some decaying leaves and twigs; full sunlight to partial shade. Cu-

lex (C. ) asteliae. |

69. Rangitoto Island, wharf (North Auckland), 7 Jan 64, BS. Cistern at

swimming pool; larvae in cistern cover and air hole. Culex (C.) quinquefasi-

atus.

~~ "0. Hooper's Inlet, sea side of road on western shore of inlet (Otago), 2

Jan 64, BN. Ditch in saltmarsh; water salty; vegetation scanty, herbaceous;

sand bottom; full sunlight. Aedes (H.) australis.

71. Hooper's Inlet, inland side of road on western shore of inlet (Otago),

2 Jan 64, BN. Ditch in cultivated field; water brackish; abundant herbaceous

vegetation and flotage; mud bottom; full sunlight. Culex (C.) pervigilans.

72. Governors Bay, Lyttelton Harbour (Canterbury), 3 Jan 64, BD. Mar-

gins of small stream; current moderate; water clear and fresh; scanty herba-

ceous vegetation, abundant flotage; rock bottom; partial to full sunlight; larvae

primarily in flotage. Nothodixa campbelli; Paradixa fuscinervis.

73A. Nelson, stream across from Gourlay's home (Nelson), 3 Jan 64, BG.

Sweeping vegetation on banks of stream (same as 73B); most abundant 1930-

2000 hrs. Paradixa neozelandica; P. fuscinervis. |

73B. Nelson, stream across from Gourlay's home (Nelson), 3 Jan 64, BG.

Margins of stream; slow to moderate current; water clear and fresh; vegeta-

tion and flotage scanty; mud bottom; partial to full sunlight. Paradixa neoze-

landica; P. fuscinervis; Culex (C.) pervigilans.

74A. North Branch Riwaka River, Crystal Pool, just below source (Nelson),

4 Jan 64, BG. Female under overhanging rock, above breeding sites (collec-

tion 74B). Paradixa fuscinervis.

74B. North Branch Riwaka River, Crystal Pool, just below source (Nelson),

4 Jan 64, BG. Stream pool; current slow; water clear and fresh; scanty her-

baceous vegetation and flotage; rock bottom; partial shade. Paradixa fusciner-

vis.

75. North Branch Riwaka River, large bridge just below fords over small

streams before source of river (Nelson), 4 Jan 64, BG. Stream margins; mod-

erate to strong current; water clear and fresh; scanty herbaceous vegetation

and flotage; gravel and rock bottom; partial shade. Nothodixa only along very

small branch of stream, not over 1 ft in width, very shallow; primarily in

shade and largely out of water. Nothodixa campbelli; Paradixa fuscinervis;

Culex (C.) pervigilans. |

76A. Canaan Beech Forest (Nelson), 4 Jan 64, BG. Adults landing in

clearing of beech forest; 1200-1300 hrs (same general location as 76B). Aedes

(O.) antipodeus. ae

76B. Canaan Beech Forest (Nelson), 4 Jan 64, BG. Tiny treeholes in

young Nothofagus, probably formed at sites of former branches of saplings;

height about 1.2 m; water very dark reddish brown, very scanty; deep shade.

Mosquitoes absent.

77. Canaan Beech Forest, Harwood's Hole Track (Nelson), 4 Jan 64, BG.

Sphagnum bog with moderately large open pond in center; water brownish;

abundant Sphagnum, Scirpus and Juncus and flotage; mud bottom; partial to

Belkin: Culicidae of New Zealand | 169

full sunlight. Culex (C.) pervigilans.

78. Canaan Beech Forest, Harwood's Hole Track (Nelson),. 4 Jan 64, BG.

Large pond with sphagnum along margins only; water a light brown; mud bot-

tom; partial to full sunlight. Culex (C.) pervigilans.

79. Aniseed Valley, Roding River, first ford below Reding ford (Nelson),

0 Jan 64, BG. Rivulet not over 2 ft in width; margins and pools; slow to mod-

erate current; fresh clear water; scanty vegetation and abundant flotage; from

deep shade to full sunlight. Larvae of Nothodixa and Paradixa together in flo-

tage in dense shade and in the open in watercress. Nothodixa campbelli; Para-

dixa fuscinervis.

80. Aniseed Valley, Roding River, third ford below Roding ford (Nelson),

o Jan 64, BG. Separate stream pools with slight flow in bed of rivulet; water

fresh and clear; scanty herbaceous vegetation and algae, abundant flotage;

sand and gravel bottom; deep shade to full sunlight. Nothodixa and Paradixa

together in all situations. Nothodixa campbelli; Paradixa neozelandica; be

fuscinervis.

81. Waitakere Ranges, West Coast Road, first stream west of Scenic

Drive (North Auckland), 8 Jan 64, BES. Stream margins; current slow to mod-

erate; water clear and fresh; vegetation and flotage scanty; gravel bottom; par-

tial to full sunlight. Paradixa fuscinervis.

82. Waitakere Ranges, Karekare Road at crossing of headwaters of Opal

Pools (North Auckland), 8 Jan 64, BES. Leaf axils of epiphytic astelias (Col-

lospermum hastatum) on blown- down tree (2 m from ground); water light brown-

ish; partial to deep shade. Culex (C.) asteliae.

83. Ohakune, Mt. Road, first stream below first bridge (Wellington), 9

Jan 64, BFS. Margins of stream dammed above culvert; moderate to strong

current; water clear, brownish; scanty fern vegetation ie flotage; mud bot-

tom; partial shade. Paradixa neozelandica: P. fuscinervis; P. harrisi.

84. Ohakune, Rangataua Reserve Lake (Wellington), 9 Jan 64, BFS. Mar-

gins of lake; water clear and light brownish; abundant herbaceous vegetation;

mud bottom; full sunlight. Paradixa fuscinervis; Culex (C.) pervigilans.

850A. Ohakune, pond on bend of Lake Road, south of Rangataua Reserve

Lake (Wellington), 9 and 10 Jan 64, BFS. Adults swept in grassy and shrubby

area around lake; very cold and windy, air temperature less than 50°F; 0815-

1015, 1700, 1930-2000 hrs. Coquillettidia (A.) tenuipalpis.

85B. Ohakune, pond on bend of Lake Road, south of Rangataua Reserve

Lake (Wellington), 9 and 10 Jan 64, BFS. Margins of lake; water cles and

light brown; abundant herbaceous vegetation (Scirpus, J nese) and ¢iotage; mud

bottom; full sunlight. Larvae primarily in clumps of vegetation. Paradixa

nébzdiandica: P. fuscinervis; Culex (C.) pervigilans.

86. Pokaka, culverts between old and new ro2ds (Wellington), 10 Jan 64,

BFS. Stream pool between 2 culverts; moderate current; water clear and

brownish; scanty herbaceous vegetation and flotage; mud pottora: full sunlight.

Paradixa neozelandica; P. fuscinervis; Culex (C. ) pervigilans.

87. Pokaka, sade: creek through pasture along main road (Wellington), 10

Jan 64, BFS. Stream margins; moderate to strong current; water clear and

brownish: scanty herbaceous vegetation and flotage; mud and sand bottom; full

sunlight. Paradixa fuscinervis.

88. Te Aroha, stream at head of main EW street (South Auckland), 10 Jan

64, BFS. Stream margin; current moderate to strong; water clear; flotage

scanty; sand and gravel bottom; deep shade. One dixine larva, apparently lost.

89. Titirangi Beach, end of Mahoe Road (North Auckland), same locality

170 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

as collections 61-63, 11 Jan 64, BS. Biting-landing in shaded area across

road from houses; 1500-1700 hrs; raining, cool. Aedes (F.) notoscriptus.

90. Titirangi Beach, end of Mahoe Road (North Auckland), same locality as -

collections 61-63, 89, 11 Jan 64, BS. Two males flying around trunk of Lepto-

spermum ericoides tree, apparently disturbed when trees looked over for rot-

holes; 1600-1700 hrs; raining. Maorigoeldia argyropus.

91. Titirangi Beach, end of Mahoe Road (North Auckland), same locality

as collections 61-63, 89,90, 11 Jan 64, BS. Paint can, tire and tin cans; water

turbid and brownish; deep to partial shade. Aedes (F.) notoscriptus.

92. Titirangi Beach, end of Mahoe Road (North Auckland), same locality

as collections 61-63, 89-91, 11 Jan 64, BS. Small rotholes in Leptospermum

ericoides, Nothopanax simplex and Pseudopanax crassifolium at height of 1-2

m above ground; water dark reddish brown; partial to deep shade. Aedes (F. )

notoscriptus.

93. Titirangi Beach, end of Mahoe Road (North Auckland), same locality

as collection 61-63, 89-92, 11 Jan 64, BS. Leaf axils of epiphytic astelias (Col-

lospermum hastatum) at 3-4 m above ground; water brownish; axils with mud;

partial shade. Culex (C.) asteliae; C.(C.) quinquefasciatus (possibly contami-

nation in laboratory).

94. Titirangi Beach, creek above dam (North Auckland), same locality as

collection 64, 11 Jan 64, BS. Margins of dammed creek; slow current; water

fresh and turbid; scanty herbaceous vegetation; abundant flotage; mud bottom;

partial shade. Paradixa neozelandica; P. fuscinervis; P. harrisi.

95. Titirangi, South Titirangi Road at beach (South Auckland), 11 Jan 64,

BS. Beached boat with small amount of brownish water; partial shade. Aedes

(F.) notoscriptus.

96. Dunedin, Botanical Gardens (Otago), 2 Jan 64, BN. Ornamental pond

with clear water; abundant herbaceous vegetation and flotage; mud bottom; par-

tial shade. Paradixa fuscinervis.

97. Waitakere Ranges, 0.25 mi S from main turnoff to Karekare from

West Coast (Piha) Road (North Auckland), 14 Jan 64, S. Roadside ditch; water

fresh and turbid; algae present; mud bottom; full sunlight. Culex (C.) pervig-

ilans.

98. Waitakere Ranges, West Coast (Piha) Road, stream east of inlet to

Nihotupu Reservoir (North Auckland), 14 Jan 64, S. Margins of stream 6-8 ft

wide; current moderate; water clear and fresh; algae present; rock bottom;

partiar-+ta deep shade. Paradixa fuscinervis.

99. Kaiapaji, Pineacres Motel (Canterbury), 21 Jan 64, S. Adults resting

inside motel room, primarily bathroom; 1900 hrs. Culex (C.) pervigilans;

Aedes (O. ) antipodeus:

100. Timaru, Scenic Réserve (Canterbury), 24 Jan 64, S. Margins of large

ground pool; water clear and fresh; abundant flotage, grassy and herbaceous

vegetation, scum and algae; mud and-decaying leaves on bottom; partial shade.

Paradixa neozelandica; Culex (C.) pervigilans.

101. Makikihi, 3 mi N of town (Canterbury), 25 Jan 64, S. Margins of

small stream; fresh, clear water; abundant grassy vegetation; mud bottom,

full sunlight. Culex (C.) pervigilans.

102. Waianakarua River, near bridge of main road (Otago), 25 Jan 64, S.

Stream margins; slow current; clear water; abundant grassy vegetation and

algae; gravel and decaying vegetation on bottom; partial shade. Paradixa neo-

zelandica; P. fuscinervis; P. tonnoiri. a

103. Clinton, 6 mi N (Otago), 26 Jan 64, S. Stream margins; moderate

Belkin: Culicidae of New Zealand ge

current; water fresh and clear; abundant grassy vegetation; partial shade. P.

fuscinervis.

104. Bluff (Southland), 26 Jan 64, S. Seaside rockholes; water clear, red-

dish and slightly salty; scanty algae; mud, gravel and rock bottom; partial to

full sunlight. Culex (C.) pervigilans; Aedes (H.) australis.

105. Oban, road to Observation Rock (Stewart Island), 27 Jan 64, S. Ditch

with muddy water; partial shade. Culex (C.) pervigilans.

106. Oban, Golden Bay at north entrance to Paterson Inlet (Stewart Island),

27 Jan 64, S. Large ground pool under rock crusher; scanty vegetation, flo-

tage, scum and algae; mud bottom with decaying vegetation; partial shade.

Culex (C.) pervigilans; Aedes (H. ) australis.

107. Between Invercargill and Bluff, just N of Awarua Radio Station (South-

land), 28 Jan 64, S. Margins of large ground pool in sheep pasture; water

fresh, turbid, brownish; scanty grassy vegetation; mud bottom; full sunlight.

Culex (C.) pervigilans.

108. Between Invercargill and Bluff, just N of fertilizer plant (Southland),

28 Jan 64, S. Flooded pasture; water fresh, turbid, brownish; abundant her-

baceous, grassy and floating vegetation, also scum and algae; mud bottom; full

sunlight. Culex (C.) pervigilans.

109A. Black Gully, 2.5 mi S of Crookston (Otago), 31 Jan 64, S. Sweeping

along stream margin; in beech forest. No mosquitoes.

109B. Black Gully, 2.5 mi S of Crookston (Otago), 31 Jan 64, S. Stream

margin; moderate to strong current; fresh, clear water; gravel bottom, par-

tial shade; in beech forest. Nothodixa campbelli: Paradixa fuscinervis.

110. Te Anau (Southland), 2 Feb 64, S. Flooded pasture; grassy vegetation;

mud bottom; full sunlight. Culex (C.) pervigilans.

111. Queenstown, Ben Lomond (Otago), 2 Feb 64, S. Stream with abundant

watercress and grass; current slow; water clear and fresh; mud bottom; par-

tial shade. Larvae clinging to watercress at water level. Paradixa neozeland-

ica; P. fuscinervis.

112. Lake Mapourika (Westland), 7 Feb 64, S. Adults biting just after dark;

weather clear. Culiseta (C.) tonnoiri; Coquillettidia (C.) iracunda.

113. Waiho vicinity (Westland), 7 Feb 64, S. Large ground pool in pasture;

water fresh and clear. Culex (C.) pervigilans.

114. Whataroa, 1 mi S of Waitangi River on main S road (Westland), 7 Feb

64, S. Large ground pool with clear fresh water and scanty vegetation; mud

and decaying vegetation on bottom; partial shade. Paradixa neozelandica; P.

fuscinervis; Culex (C.) pervigilans.

115. Hercules Mt vicinity, 7-8 mi S of Harihari on route 6 (Westland), 7

Feb 64, S. Margin of stream pool; slow current; water clear; scanty grassy

vegetation, flotage, scum and algae present; mud bottom; full sunlight. Para-

dixa neozelandica; P. fuscinervis; P. tonnoiri; Culex (C.) pervigilans.

116. New Creek Rd, between Murchison and Kawatiri Junction (Nelson), 8

Feb 64, S. Stream pool; slow current; scanty grassy yegetation; sand and

rock bottom; partial shade. Nothodixa campbelli; Paradixa neozelandica; P.

fuscinervis; Culex (C.) pervigilans. :

117A. Lake Rotoiti, very small stream above campground (Nelson), 9 Feb

64, S. Sweeping in virgin Nothofagus forest; 1100 hrs; clear and sunny. No

mosquitoes.

117B. Lake Rotoiti, very small stream above campground (Nelson), 9 Feb

64, S. Stream margin; slow current; water clear and fresh; scanty grassy

vegetation; gravel and mud bottom; partial to deep shade. Paradixa fusciner-

vis.

172 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

118. Lake Rotoiti, campground (Nelson), 9 Feb 64, S. Small treehole in

Nothofagus; height 1-2 m; reddish brown water with decaying vegetation on

bottom; partial shade. No mosquitoes.

119. Lake Rotoiti, above campground (Nelson), 9 Feb 64, S. Large tree-

hole in Nothofagus; height 0.3 m; greenish-yellow water; decaying vegetation

on bottom; partial to deep shade. Maorigoeldia argyropus.

120. Kawatiri Junction, junction of Buller and Hope roads (Nelson), 9 Feb

64, S. Large ground pool; water clear; abundant herbaceous and grassy vege-

tation; gravel bottom; full sunlight. Culex (C.) pervigilans.

121. Gowan River on way to Lake Rotorua (Nelson), 9 Feb 64, S. Stream

margin; Slow current; water clear; abundant grass and watercress, also algae;

mud bottom; partial shade. Paradixa neozelandica; Culex (C.) pervigilans.

122. Gowanbridge, near junction of Buller and Gowan rivers (Nelson), 9

Feb 64, S. Large ground pool in seepage area; slow.current; water fresh and

clear; abundant watercress; mud bottom; partial shade. Paradixa neozelandi-

ca; P. fuscinervis; Culex (C.) pervigilans.

123A. Nelson, North Branch Maitai River (Nelson), elev. ca130 m, 11 Feb

64, S. Sweeping along banks of small stream and resting on Nothofagus trunk.

Maorigoeldia argyropus. |

123B. Nelson, North Branch Maitai River (Nelson), elev. ca130 m, 11 Feb

64, S. Margin of stream; slow current; fresh, clear water; vegetation very

scanty; gravel and rock bottom; deep shade. Paradixa fuscinervis.

124. Nelson, Cawthron Park (Nelson), elev. ca 150 m, 11 Feb 64, S. Small

ground pool in seepage; slow current; clear, fresh water; scanty vegetation;

mud and decaying vegetation on bottom; partial to deep shade. Paradixa fusci-

nervis.

125. Nelson, junction of Maitai River and Sclanders stream (Nelson), 11

Feb 64, S. Stream margin and stream pools; slow current; clear, fresh water;

scanty vegetation, scum and algae; mud and decaying vegetation on bottom;

partial shade in Nothofagus forest. Nothodixa campbelli; Paradixa fuscinervis;

Culex (C. ) pervigilans.

126. Nelson, South Branch Maitai River (Nelson), 11 Feb 64, S. Small

ground pool in seepage; scum present. Culex (C.) pervigilans.

127. Nelson, South Branch Maitai River (Nelson), 11 Feb 64, S. Margins

of small stream flowing into river; scanty watercress and grassy vegetation;

mud and sand bottom; partial shade. Nothodixa campbelli; Paradixa fusciner-

vis. We eel reine oka One Sig

~- 128. Little Barrier Island, small stream between Te Waikohare and Tiriki-

kawa streams (Hauraki Gulf), elev. 7m, 6 Mar 64, S. Separate pools in almost

dry stream bed. Nothodixa campbelli; Culex (C.) pervigilans; Aedes (O.) an-

tipodeus; Aedes (F.) notoscriptus; Maorigoeldia argyropus.

129. Little Barrier Island, near chicken house (Hauraki Gulf), elev. 7 m, 6

Mar 64, S. Small artificial container with brownish fresh water. Culex (C.)

pervigilans; Aedes (F. ) notoscriptus.

130. Little Barrier Island, Waipawa stream, 75-100 ft from mouth (North

Auckland), 6 Mar 64, S. Separate pools in dry stream bed; clear stagnant wa-

ter; scanty grassy vegetation and flotage; bottom with decaying leaves; partial

shade. Paradixa neozelandica; P. fuscinervis; P. harrisi; Culex (C.) pervig-

ilans; Aedes (F.) notoscriptus.

131. Little Barrier Island, Tirikikawa stream (Hauraki Gulf), elev. 30 m,

8 Mar 64, S. Separate pools in nearly dry stream bed; slow current; fresh

clear water; some flotage; decaying vegetation on bottom; partial shade. Pa-

radixa fuscinervis; P. harrisi; Culex (C.) pervigilans.

Belkin: Culicidae of New Zealand 173

132. Little Barrier Island, Summit Track (Hauraki Gulf), elev. 225 m, 7

Mar 64, S. Leaf axils of terrestrial astelias (?Collospermum hastatum). Cu-

lex (C. asteliae.

133. Little Barrier island. Summit Track (Hauraki Gulf), elev. 250 m, 7

Mar 64, S. Treehole in Nothofagus, 2x 3in; water brownish; in virgin forest.

Maorigoeldia argyropus.

134. Little Barrier Island, Hut Bay Creek (Hauraki Gulf), near sea level,

8 Mar 64, S. Stream margins and pools; slow current; clear water; some flo-

tage; decaying vegetation on bottom; partial shade. Paradixa neozelandica;

P. fuscinervis; P. harrisi.

135. Mokohinau Island (Hauraki Gulf), on shipboard, 11 Mar 64, S. Fe-

males on human bait; 1630 hrs; windy. Island mostly cleared, numerous rock-

pools. Opifex fuscus.

136. Cuvier Island (Hauraki Gulf), 13 Mar 64, S. Ground pool. Culex (C.)

pervigilans.

137. Cuvier Island, 13 Mar 64, S. Seaside volcanic rockhole; water brack-

ish; mud bottom; partial shade. Opifex fuscus.

138. Great Barrier Island, Port Fitzroy (Hauraki Gulf), 12 Mar 64, S.

Stream margins. Nothodixa campbelli; Paradixa neozelandica; P. fuscinervis;

P. harrisi.

139. Te Aroha Stoney Creek (South Auckland), 26 Mar 64, 8. Stream mar-

gin; Slow to moderate current; clear, fresh water; no vegetation or flotage;

gravel and rock bottom; partial shade. Nothodixa campbelli: N. septentrional-

is.

140. Ormond, Waihirere Domain (Gisborne), 28 Mar 64, S. Stream mar-

gins and pools below falls; slow current; clear, fresh water; scanty grassy

vegetation; partial shade. Paradixa neozelandica; P. fuscinervis; Culex (C.)

pervigilans.

141. Urewera National Park, Hopuruahine area, Orangihikoa Stream (Gis-

borne), elev. 2450 ft, 30 Mar 64, S. Stream margin; moderate current; clear,

fresh water; scanty grassy vegetation. Nothodixa campbelli; N. septentrional-

is; Paradixa fuscinervis.

142. Te Puke, Prew Orchard, #3 road (South Auckland), 31 Mar 64, 8. Ce-

ment water basin, 3 ft diameter, 1 ft deep; bottom with decaying pine needles;

partial shade. Culex 4 One) pervigilans ; Aedes (F.) notoscriptus.

143. Clevedon (North Auckland), 5 Apr 64, S. Metal cooking pan, 1 ft sq,

3 in high; rusty water; decaying leaves and twigs on bottom; partial shade in

native bush. Maorigoeldia argyropus.

144. Clevedon (North Auckland), 5 Apr 64, S. Tire; mud and decaying veg-

etation on bottom; partial to deep shade in native bush. Maorigoeldia argyro-

us.

145-149. No collections.

150. Chatham Island, Te Awarakau near Waitangi, elev. 4m, 20 Feb, 8.

Large seaside rockpools and seepages; slow current; water fresh to salty;

scanty flotage and algae; sand and rock bottom with decaying vegetation; full

sun. See also 155. Aedes (N.) chathamicus.

151. Chatham Island, road near flying boat base, 26 Feb 64, S. Large

ground pool; yellowish fresh water; abundant flotage and algae; mud bottom;

full sun. Culex (C.) pervigilans.

152. Chatham Island, road just S of Owenga, 26 Feb 64, S. Stream mar-

gin; slow current; muddy water; abundant grassy vegetation; mud bottom; full

sun. Culex (C.) pervigilans.

174 Contrib. Amer. Ent..Inst., vol: 3,.no. 1, 1968

153. Chatham Island, Waitangi West, 28 Feb 64, S. Large ground pool;

clear fresh water; scanty algae; mud and sand bottom; full sun. Culex (C.).

pervigilans.

154. Chatham Island, Waitangi, 29 Feb 64, S. Clear rainwater in dinghy;

partial shade. Culex (C.) pervigilans.

155. Chatham Island, Te Awarakau near Waitangi, elev. 4 m, same locali-

ty as collection 150, 2 Mar 64, S. Large seaside rockpools and seepages;

slow current; water fresh to salty; scanty flotage and algae; sand and rock

bottom with decaying vegetation; full sun. Aedes (N.) chathamicus.

156. Springs Junction, Springsland Creek behind Rahu Post Office (Nelson),

elev. 1820 ft, 9 Mar 64, S. Stream margins; moderate current; clear, fresh

water; scanty grassy vegetation; gravel bottom; full sun. Paradixa fusciner-

vis. Co (rec erce,

157. Tahunanui, stream E of Motor Camp (Nelson), elev. 3 m, 10 Mar 64,

S. Stream margin; moderate current; fresh water, muddy due to flooding;

abundant grassy vegetation; mud bottom; partial shade. Culex (C.) pervigilans;

Aedes (O. ) antipodeus.

158. Annesbrook (Nelson), elev. ca20 m, 10 Mar 64, S. Stream margin;

moderate current; clear water; scanty grassy vegetation; gravel bottom; par-

tial to full sun. Paradixa neozelandica; P. fuscinervis.

159. Ngawhatu Mental Hospital, streams along road (Nelson), 11 Mar 64,

S. Stream margins; moderate current; clear, fresh water; grassy vegetation,,

partial shade. Paradixa fuscinervis.

160. Stoke, Polstead Road (Nelson), 11 Mar 64, S. Swamp margin; stag-

nant, clear, fresh water; abundant grassy vegetation. Culex (C.) pervigilans.

161. Brightwater, main highway bridge over Waimea River (Nelson), 11

Mar 64, S. River margin; moderate current; turbid fresh water; scanty

grassy vegetation; mud, sand and gravel bottom; partial shade. Paradixa

fuscinervis.

162. Nelson, Reservoir (Nelson), 12 Mar 64, S. Stream margin; slow cur-

rent; clear, fresh water; scanty vegetation; mud, sand, gravel and rock bot-

tom; deep shade. Paradixa neozelandica; P. fuscinervis.

163. Nelson, Reservoir (Nelson), 12 Mar 64, S. Tire with mud and decay-

ing vegetation on bottom; partial shade. Culex (C.) pervigilans.

164. Blick's Valley, stream off Brook Stream (Nelson), 12 Mar 64, S.

Stream margin; slow to moderate current; clear fresh water; grassy vegeta-

tion; partial to full sun. Paradixa neozelandica; P. fuscinervis; Culex (C.)

pervigilans.

165. Dodson's Valley Stream (Nelson), elev. ca68 m, 12 Mar 64, S. Stream

margin; Slow to moderate current; clear fresh water; abundant grassy vege-

tation; mud and gravel bottom; partial shade. Nothodixa campbelli; Paradixa

neozelandica; P. fuscinervis.

166. Todd's Valley Stream (Nelson), 12 Mar 64, S. Stream margin; slow

to moderate current; clear fresh water; grassy vegetation; mud and gravel

bottom. Paradixa neozelandica; P. fuscinervis.

167. Wakapuaka, Glen Road (Nelson), 12 Mar 64, 8S. Ditch under road;

slow current; clear fresh water; grassy vegetation; mud bottom; partial

shade. Paradixa neozelandica; P. fuscinervis; Culex (C.) pervigilans.

168. The Glen (Nelson), 12 Mar 64, S. Stream margin just before beach;

slow current; clear fresh water; grassy vegetation; gravel and rock bottom.

Nothodixa campbelli; Paradixa neozelandica; P. fuscinervis.

169. Howard Junction, 1 mi E, along Buller River (Nelson), 13 Mar 64, S.

Belkin: Culicidae of New Zealand 175

Large ground pool adjacent to river; clear fresh water; abundant grassy veg-

etation and duckweed; partial shade. Paradixa neozelandica; P. fuscinervis;

P. tonnoiri; Culex (C.) pervigilans.

170. Nelson, Maitai River, near Pole Ford Bridge (Nelson), 14 Mar 64, S.

Ditch along road; slow current; clear fresh water; mud bottom; partial shade.

Paradixa neozelandica; P. fuscinervis; Culex (C.) pervigilans.

171. Nelson, Maitai River and Dod's Creek at Smith's Ford (Nelson), 14

Mar 64, S. Stream margin; slow current; clear fresh water; mud and gravel

bottom; partial shade. Paradixa neozelandica; P. fuscinervis.

172. Rakino Island (Hauraki Gulf), 25 Jan 64, F. Seaside rockpool above

high tide; chloride 34000 ppm. Opifex fuscus.

173. Rakino Island (Hauraki Gulf), 24 Jan 64, F. Seaside rockpool above

oyster zone; chloride 34000 ppm. Opifex fuscus.

174. Ohakune, pond on bend of Lake Road, south of Rangataua Reserve,

Lake (Wellington), 26 Jan 64, F. Same locality as collection 85. Culex (C.)

pervigilans.

175. Ohakune, Mt. Road, first stream below first bridge (Wellington), 26

Jan 64, F. Same locality as collection 83. Paradixa fuscinervis.

176. Ohakune, pond on bend of Lake Road, south of Rangataua Reserve Lake

(Wellington), 28 Jan 64, F. Same locality as collections 85 and 174. Culex (C.)

pervigilans.

177. Waitakere Ranges, Cascade Park, Waitakere stream (North Auckland),

20 Feb 64, F. Sweeping and netting. Paradixa fuscinervis; Coquillettidia (C. )

iracunda.

178. Waitakere Ranges, Cascade Park, Waitakere stream, bridge below

Cascade Falls (North Auckland), 20 Feb 64, F. Paradixa fuscinervis.

179. Waitakere Ranges, Bethells Road near Quarry (North Auckland), 11

Apr 64, F(D1). Stream margins and pools; slow current; clear fresh water;

no vegetation; rock and mud bottom; partial shade. Paradixa fuscinervis; P.

harrisi.

180. Mangere Sewage Plant (North Auckland), 22 Apr 64, F(D3). No. 1 oxi-

dation pond; water green with algae; mud bottom; full sun. Culex (C.) quin-

quefasciatus.

181. Henderson, Opunaku Stream (North Auckland), elev. 65 m, 26 Apr 64,

F(D4). Stream pool; slow current; clear fresh water; some Potamageton;

mud and rock bottom; partial shade. Paradixa fuscinervis.

182. Albany, Rosedale Road (North Auckland), elev. 100 m, 18 May 64, F

(D5). Stream pool; slow current; clear fresh water; mud and rock bottom

with decaying vegetation; deep shade. Paradixa neozelandica; P. fuscinervis.

183. Swanson, Train Valley Stream (North Auckland), elev. 166 m, 9 May

64, F(D6). Stream pool; slow current; turbid yellowish water; partial shade.

Paradixa fuscinervis.

184. Waitakere Ranges, Bethells Road Swamp (North Auckland), 9 May 64,

F(D7). Swamp margin; yellowish-brown fresh water; abundant vegetation of

Typha and reeds; mud bottom with decaying vegetation; partial shade; 2 in rain

in previous 5 days. Culex (C.) pervigilans.

185. Rotorua, Whakarewarewa (South Auckland), 19 May 64, F(D8). Mar-

gins of stream bounding thermal area; temperature of water normal not ther-

mal; slow current; turbid water with slight sulfurous odor; scanty vegetation;

sand bottom; partial shade. Culex (C.) rotoruae.

186. Pukehoke, Sewage Oxidation Pond (North Auckland), elev. 60 m, 25

May 64, F(D9). Oxidation pond; clear fresh temporary water; scanty grassy

176 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

and herbaceous vegetation; mud bottom; full sun. Culex (C.) pervigilans.

187A. Ocean Beach, Whangarei Heads (North Auckland), 31 May 64, F(D10).

Landing and resting collection of adults at rockpool; 1445 hrs; clear, sunny.

Opifex fuscus.

187B. Ocean Beach, Whangarei Heads (North Auckland), 31 May 64, F(D10).

Seaside volcanic rockholes at 9 ft above high tide level; clear water; sand bot-

tom; full sun. Opifex fuscus.

188. Wellsford, stream under route 1 north of town (North Auckland), elev.

100 m, 31 May 64, F(D11). Stream margin; slow current; clear fresh water;

sparse growth of Typha; mud bottom; partial shade. Paradixa fuscinervis;

Culex (C.) pervigilans.

189A. Titirangi, French Bay (North Auckland), elev. 50 m, 14 June 64, F

(D12). Dixids flying over breeding site 189B; 1200 hrs; showery and humid.

None collected.

189B. Titirangi, French Bay (North Auckland), elev. 50 m, 14 June 64, F

(D12). Stream pool; slow current; turbid fresh water; scum; mud bottom;

deep shade. Paradixa fuscinervis; P. harrisi.

190. Tuakau, bridge over Waikato River (North Auckland), elev. ca 10m, 9

June 64, F(D13). Ditch into Waikato River; slow current; clear fresh water;

no vegetation; mud bottom; partial shade. Paradixa fuscinervis; Culex (C.)

pervigilans.

191A. Titirangi Beach, Mahoe Road (North Auckland), elev. 6 m, 4 July 64,

F(D14). Landing on collector: 1100 hrs. Aedes (F.) notoscriptus.

191B. Titirangi Beach, Mahoe Road (North Auckland), elev. 6 m, 4 July 64,

F(D14). Small tin can with clear water; oil on surface; mud bottom; deep

shade. Aedes (F.) notoscriptus; Maorigoeldia argyropus.

192A. Titirangi, French Bay (North Auckland), elev. 5 m, 4 July 64, F(D15).

Adults swarming over ditch, about 1m; 1100 hrs. Paradixa neozelandica; P.

fuscinervis. |

192B. Titirangi, French Bay (North Auckland), elev. 5 m, 4 July 64, F(D15).

Ditch; slow current; turbid fresh water; flotage; mud bottom with decaying

vegetation; deep shade. Paradixa neozelandica; P. fuscinervis.

193. Titirangi Beach, Mahoe Road (North Auckland), elev. 2m, 4 July 64,

F(D16). Stream margin; slow current; turbid fresh water; no vegetation; mud

bottom; partial shade. Paradixa fuscinervis. -

194. Avondale, Auckland, Methuen Road (North Auckland), elev. 60m, 9

Aug 64, F(D15!). Washing machine bowl in domestic yard; yellowish water

with decomposing leaves; deep shade. Culex (C.) quinquefasciatus; Aedes (F. )

notoscriptus.

195A. Waitakere Ranges, on Bethells Road ca10 yd from Black bridge and

bridge on Cascade turnoff (North Auckland), elev. 20 m, 15 Nov 64, F(D17).

Sweeping collection.

195B. Waitakere Ranges, on Bethells Road ca10 yd from Black bridge and

bridge on Cascade turnoff (North Auckland), elev. 20 m, 15 Nov 64, F(D17).

Ground pool; yellowish, fresh water; mud bottom with decaying vegetation;

partial shade. Paradixa neozelandica; P. fuscinervis.

195C. Titirangi Beach, Mahoe Road (North Auckland), elev. 5 m, 29 Nov 64,

F. Coquillettidia (C.) iracunda.

196. Martins Bay, Mahurangi Heads (North Auckland), elev. 3 m, 26 Dec

64, F(D21). Marshy depression in swamp; turbid fresh water; abundant wa-

tercress: mud bottom; full sun. Paradixa neozelandica; Culex (C. ) pervigilans.

197. Martins Bay, Mahurangi Heads (North Auckland), 29 Dec 64, F(D23).

Belkin: Culicidae of New Zealand 177

Blocked ditch on beach; clear fresh water; abundant scum and sedges; mud

bottom with decaying vegetation; partial shade. Paradixa neozelandica.

198A. Mangawhai Heads, Waipu Road (North Auckland), elev. 12 m, 2 Jan

65 F(D26). Stream margin; Slow current; clear fresh water; flotage pres-

ent, no vegetation; gravel bottom; partial shade. Paradixa fuscinervis.

198B. Dome Valley (North Auckland), 2 Jan 65, F. Paradixa neozelandica;

P. fuscinervis.

199. Piriaka, 6 mi S of Taumarunui (Wellington), elev. 200 m, 5 Jan 65,

F(D28). Stream pool in tributary of Wanganui River, 4 x 8 ft, 10 in deep; slow

current; clear fresh water; flotage of willow leaves; mud bottom with willow

roots; partial shade. Paradixa neozelandica; P. fuscinervis.

200A. Waitakere Ranges, Bethells Road Swamp (North Auckland), 10 Jan

65, F(D29). Biting and sweeping collection, 1630 hrs. Coquillettidia (C.) ira-

cunda; Aedes (F. ) notoscriptus.

200B. Waitakere Ranges, Bethells Road Swamp (North Auckland), 10 Jan

65, F(D30). Swamp margin; reddish brown fresh water; abundant Typha and

reeds; deep shade. Paradixa neozelandica; P. fuscinervis; Culex (C.) pervig-

ilans; Coquillettidia (A. ) tenuipalpis; C.(C.) iracunda.

201. Mangere (North Auckland), 7 Jan 65, F(D31). Culex (C.) pervigilans.

202. Waitakere Ranges, Bethells Road Swamp (North Auckland), 12 Jan 65,

F(D32). Same locality and data as collection 200B. Paradixa neozelandica; P.

fuscinervis; Culex (C.) pervigilans.

203A. Dome Valley, southern end, near Warkworth (North Auckland), 16

Feb 65, F(D32). Sweeping under bridge; 1000 hrs. Dixids.

203B. Dome Valley, southern end, near Warkworth (North Auckland), 16

Feb 65, F(D32). Stream margin; slow current; turbid, reddish fresh water;

grassy vegetation; mud bottom; deep shade. Culex (C.) pervigilans.

204A. Wellsford, first bridge S of city on highway 1 (North Auckland), 16

Feb 65, F(D33). Stream pool; clear fresh water; abundant grassy and herba-

ceous vegetation; mud bottom with decaying vegetation; deep shade. Paradixa

fuscinervis.

204B. Kaitaia, Kaitaia Hotel (North Auckland), 16 Feb 65, F. On wall in

hotel. Coquillettidia (C.) iracunda.

205. Mangamuka Gorge, E side of highway 1, dammed section of stream

marked "Fresh Water" (North Auckland), elev. ca500 m, 18 Feb 65, F(D35).

Stream margin and pool; slow current; clear fresh water; submerged vege-

tation, Callitriche, Nitella; mud bottom; deep shade. Nothodixa campbelli;

Paradixa harrisi.

206. Mangamuka Gorge, S slope, at bridge (North Auckland), 18 Feb 65,

F(D36 and D36A). Stream pool; slow current; clear fresh water; scanty veg-

etation; mud bottom; deep shade. Paradixa fuscinervis; P. harrisi.

207. Brynderwyn Roadside, S slope (North Auckland), elev. ca 300 m, 18

Feb 65, F(D37). Stream pool at foot of waterfall; slow current; clear fresh

water; scanty flotage and vegetation, Callitriche; mud and gravel bottom; par-

tial shade. Paradixa harrisi.

208. Dome Valley, bridge S of Wellsford (North Auckland), 18 Feb 65, F

(D38). Stream margin and pool; slow current; turbid fresh water; some flo-

tage; mud bottom with decaying vegetable debris; deep shade. Paradixa fusci-

nervis. |

209A. Mangere Sewage Treatment Plant (North Auckland), 9 Mar65, F(D40).

Adults swept from grassy margin of pond. Paradixa neozelandica.

209B. Mangere Sewage Treatment Plant (North Auckland), 9 Mar 65, F(D40).

178 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

Pond; turbid, yellowish fresh water; abundant algae and grassy vegetation;

mud bottom; full sun. Paradixa neozelandica.

209C. Mangere Sewage Treatment Plant (North Auckland), laboratory colo-

ny established from collection 209B, 9 Mar 65, F(D40,D42). Paradixa neoze-

landica.

210. Piriaka, S of Taumarunui (Wellington), elev. 200 m, 28 Mar 65, F

(D43). Stream margin; slow current; clear fresh water; abundant grassy

vegetation; mud bottom with decaying vegetation; partial shade. Paradixa neo-

zelandica; P. fuscinervis.

211. Te Aroha, 10 miS, ca0.5 mi from waterfall on dead end road (South

Auckland), 5 June 65, F(D45). Stream margin; slow current; clear fresh wa-

ter; abundant flotage and grassy vegetation; mud and sand bottom with decaying

vegetation; partial shade. Paradixa neozelandica; P. fuscinervis; P. harrisi.

212A. Ohakune, Rangataua Reserve Lake (Wellington), elev. 600 m, 6 June

65, F(D46). Sweeping along margin of lake; 1620 hrs. Paradixa neozelandica.

212B. Ohakune, Rangataua Reserve Lake (Wellington), elev. 600 m, 6 June

65, F(D46). Lake margin; clear fresh water; abundant vegetation; mud bot-

tom with decaying vegetation; partial shade. Paradixa neozelandica.

213. Between Piriaka and Taumarunui, on highway 4 at Manunui turnoff

(Wellington), 7 June 65, F(D47). Ditch; slow current; turbid, brownish fresh

water; abundant pasture grass; mud bottom with decaying vegetation; partial

shade. Paradixa neozelandica; P. fuscinervis.

214. Mapiu, picnic area ca5 mi S on highway 4 (South Auckland), 7 June 65,

F(D48). Ditch; moderate current; clear fresh water; abundant vegetation;

mud bottom with decaying vegetation; partial shade. Paradixa fuscinervis.

215. Mangere Purification Plant, experimental pond A (North Auckland), 15

June 65, F(D49). Pond; turbid, yellowish fresh water; abundant flotage, al-

gae and grassy vegetation; mud bottom with decaying pasture grass; full sun.

Paradixa neozelandica.

216A. Mangere Purification Plant, experimental pond B (North Auckland),

15 June 65, F(D50).. Pond; turbid, yellowish fresh water; abundant flotage,

algae and pasture grass; mud bottom with decaying vegetation; full sun. Para-

dixa neozelandica.

216B. Avondale (North Auckland), elev. 60 m, 23 Nov 64-11 Jan 65, F.

Paradixa neozelandica; P. fuscinervis; Culex (C.) pervigilans.

217. Blackhead, Dunedin (Otago), 23 Feb 64, N. Stream margin about 100

yd from shore; very slow current; watercress. Paradixa neozelandica.

218. Coromandel (South Auckland), 21 Apr 64, N. Boat hull. Culex (C.)

quinquefasciatus.

219. Coromandel (South Auckland), 22 Apr 64, N. Astelia axils. Culex (C.)

asteliae.

220. Coromandel (South Auckland), 25 Apr 64, N. Weedy ditch. Culex (C.)

pervigilans. |

221. Coromandel (South Auckland), 25 Apr 64, N. Overgrown mine work-

ings. Paradixa neozelandica.

222. Coromandel (South Auckland), 26 Apr 64, N. Domestic water tank.

Aedes (F. ) notoscriptus.

223. Hooper's Inlet, Dunedin (Otago), 3 Aug 63, N. Ditch. Aedes (O.) sub-

albirostris.

224. Bridge Flat-Haast Road, NZ S1 (Westland), 18 Dec 66, N. Slow-flow-

ing water on forest edge. Paradixa fuscinervis; P. tonnoiri.

225. Waitakere Ranges, Bethells Road Syanp (North Auckland), 6 Dec 66,

D. Coquillettidia (A.) tenuipalpis.

Belkin: Culicidae of New Zealand 179

APPENDIX B

Laboratory Colonization of Paradixa neozelandica Tonnoir

Don Forsyth!

Colonies of the dixa midge Paradixa neozelandica (Tonnoir, 1924) can be

readily established and maintained in the laboratory for indefinite periods un-

der quite simple conditions.

Due to the wide degree of individual variation in features of the wing vena-

tion and male genitalia observed in adults of this species collected throughout

the Auckland province, New Zealand, colonization under artificial conditions

was valuable as a means of determining the range of variation and establishing

whether or not a character bred true.

The breeding unit comprised a clear plastic tank 16 inches by 10 inches by

8 inches deep, on the floor of which was placed a "'Hykro" underground filter,

type P 70, size 2. The vertical air outlet pipe of this was cut down to 5 inches

and the offcut cemented to the basal piece at a right angle and parallel to the

short sides of the tank. The water level was taken to 1/2 inch above this hori-

zontal limb and the inlet air pipe connected to an aerator. The entry of air was

adjusted to give a moderate flow of water at a rate of 2 feet per second which

was deflected from the opposite wall to circulate around the tank thus simu-

lating conditions encountered in the field. It was found necessary to place some

weights upon the filter to prevent it from floating and to keep it in a fixed posi-

tion on the floor of the tank since no substrate material was used as this would

defeat the purpose of the filter, i.e. that of keeping the solid organic particles

in the water in suspension and circulation (the orthodox use of an underground

filter being to remove such materials from suspension and to maintain the wa-

ter ina clear and well-aerated condition).

Three feeding methods were used to maintain the culture:

1) Water from the larval collecting site along with a small amount of sub-

strate to provide natural nutrients.

2) Tap water into which soya bean flour was introduced daily, rapidly pro-

ducing a flourishing growth of assorted unicellular algae and protozoans.

3) Tap water with the addition of a pure culture of the unicellular green al-

ga, Euglena sp.

Of these, the second method proved to be most effective in that it was sim-

ply maintained and produced the shortest life cycle, 20 days from egg to adult.

However, in some areas the local tap water may not be entirely suitable owing

to the method of pre-treatment and in such cases methods (1) and (2) could be

combined.

The colony was established from 24 fourth-instar larvae collected in the

field and introduced into the tank which was covered by fine muslin. The lar-

vae pupated and adults subsequently emerged, some of the females producing

IDepartment of Zoology, University of Bristol, Bristol, England.

180 Contrib. Amer. Ent. Inst., vol. 3, no. 1, 1968

fertile eggs. The latter were enclosed ina circular, clear, gelatinous matrix

(25 to 78 eggs per mass) and were white at first and later became dark brown

as development proceeded. The egg masses were deposited on the tank walls

at the meniscus directly opposite the water outlet where the flow was greatest,

regardless of light intensity or direction.

Each female laid a single clutch of eggs which hatched with an average pe-

riod of time of 4 days. The young first instar larvae remained within the gelat-

inous matrix for up to 24 hours before swimming freely when they were swept

away to less turbulent parts of the tank where they pushed themselves upwards

on the tank walls out of the meniscus to assume their characteristic inverted

U-shaped attitude within a thin film of water.

Even as later generations became out of phase and overlapped and larval

density rose, the second, third and fourth-instar larvae continued to inhabit the

less turbulent areas so that the newly hatched larvae while still within the egg

mass were not subjected to their interference. :

For the 3 methods used the life cycle averaged 30 days at 20°C. Adult

life averaged 4 days during which time no food was apparently taken. The fe-

males oviposited within 72 hours of emergence.

The colony was maintained continuously for 5 generations (before being dis-

carded) without larval mortality although 30 percent of the egg masses produced

did not hatch, probably because they were not fertilized.

This is the first record of the laboratory colonization of a dixa midge and

it is to be hoped that the method can be adopted by future workers for other

members of the group.

INDEX TO SCIENTIFIC NAMES

absorbrina (Culiseta, Culex) 71 asteliae (Culex) 5, 50, 54 & 55 (keys),

*acer (Culex) 68 58, 60, 65, 66, 134 & 135 (figs.),

Aedes 51 & 52 (keys), 82 & 83 & 84 152 (map), 163-165, 168-170, 173,

(keys), 90 178.

Aedini 80 astelias 164, 165, 166, 167, 173, 178

aegypti (Aedes) 10 australicus (Culex pipiens) 54

albirostris (Aedes, Ochlerotatus) 102 | australis (Aedes, Halaedes, Culex,

annulata (Culiseta, Theobaldia) 71 Pseudoskusea) 5, 6, 50, 51, 81,

annulirostris (Culex) 113 82 & 83 & 84 (keys), 86-88, 92,

anomalus (Neochaoborus) 17 93, 94, 96, 97, 157 & 158 (maps),

antipodea (Culiseta) 73 168, 171

antipodeus (Aedes, Ochlerotatus) 81, | australis (Cordyline) 166

82 & 83 & 84 (keys), 98, 102, 159 Austromansonia 50, 76

(map), 163-165, 168, 170, 172, Avicennia 165 yt

174

argyropus (Maorigoeldia, Culex, brakeleyi (Corethrella, Corethra) 110

Tripteroides, Rachionotomyia,

Uranotaenia) 5, 106, 144 (figs.), *Caenocephalus 94

161 (map), 167, 170, 172, 173, 176] caledonicus group 108

*ashworthi (Aedes, Ochlerotatus, californica (Paradixa) 10

Pseudoskusea) 94 Callitriche 177

Astelia 68

INDEX TO SCIENTIFIC NAMES

campbelli (Nothodixa, Dixa) 3, 17, 20,

21 & 22 (keys), 26, 29, 37, 41, 48,

49, 124 & 125 (figs.), 145 (map),

165, 168, 169, 171-174, 177.

Chadoborinae 7 & 8 (keys), 110

chathamicus (Aedes, Nothoskusea) 5,

6, 60, 81, 82 & 83 & 84 (keys), 86,

87, 90, 95, 139 & 140 & 141 (figs.),

157 & 158 (map), 173, 174

chilensis (Nothodixa) 10

Climacura (Culiseta) 50, 71

*concolor (Caenocephalus, Aedes,

-Pseudoskusea) 94

confirmatus (Aedes, Ochlerotatus) 98

Coquillettidia 50, 51 & 52 (keys), 74

Cordyline 67 fon

Corethrella 110

Corethrellini 110

crassifolium (Pseudopanax) 170

crassipes group (Coquillettidia) 75

*crucians (Culex, Aedes, Pseudosku-

sea) 94

Culex 50, 51 & 52 & 53 (keys), 53,

Culicinae 49

Culicini 53

Culiseta 51 & 52 & 53 (keys), 71, 76

Culisetini 71

Dixa 14, 17

Dixapuella 14

Dixella 14

*Dixella 14, 16, 29

Dixina 14

Dixinae 7 & 8 (keys)

Dixini 15 (keys)

ensifera (Nothodixa) 10

ericoides (Leptospermum) 166, 170

Erioptera 14

Eriopterites 14

*fatigans (Culex, Culex pipiens) 68

Finlaya 82 & 83 & 84 (keys), 103

fusca (Nothofagus) 109

fuscinervis (Paradixa, Dixa) 3, 15,

24, 32, 33 & 34 (keys), 37, 38, 39,

46, 49, 65, 127 & 130 (figs.), 148

(map), 163, 165-178

fuscus (Opifex) 60, 80, 81, 82 & 83 &

84 (keys), 122 & 138 (figs.), 157 &

158 (maps), 164, 167, 173, 175,

176

181

globocoxitus (Culex) 54

Halaedes 50, 51, 82 & 83, & 84 (keys),

harrisi (Paradixa, Dixa) 10, 15, 32,

33, 34 (keys), 36,37, 46, 48, 127&

132 (figs.), 149 (map), 169, 170,

172, 175-178

hastatum (Collospermum) 67, 163,

168, 169

humeralis (Paradixa) 32

inepta (Corethrella, Ramcia) 110

inexpectatus (Aedes, Ochlerotatus) 81,

iracunda (Coquillettidia, Culex, Man-

sonia, Taeniorhynchus) 3, 74, 75

(keys), 76, 78, 137 (figs.), 156

(map), 165, 167, 171, 175-177

iracunda group 76

iyengari (Culex) 53, 54

Juncus 167, 168, 169

kochi (Aedes, Finlaya, Culex) 103

linealis (Coquillettidia) 75, 76

lirio (Dixella) 31, 32

*Lutzomiops 110

maculipennis (Anopheles) 113

Mansoniini 74

Maorigoeldia 50, 51 & 52 (keys), 82,

106

marginata (Dixapuella) 31

melanura (Culiseta, Climacura,

Culex) 71, 73

Meringodixa 14

minuta (Neodixa, Dixella) 3, 16, 37,

146 (map)

miraculosus (Culex) 53, 81

molestus (Culex) 54

‘‘molestus’’ (Culex) 54

Neodixa 14, 15 (keys), 16

neozelandica (Paradixa, Dixa) 15, 32,

33, 34 (keys), 35, 41, 45, 46, 48,

49, 121 & 128 & 129 (figs.), 147

(map), 165-178, 179

nicholsoni (Paradixa) 32

nigra (Corethrella, Lutzomiops) 110

nikau palm 67

Nitella 177

182 INDEX TO SCIENTIFIC NAMES

nitida (Nothodixa) 10

Nothodixa 14, 15 (keys), 17

Nothofagus 168, 171, 172, 173

Nothoskusea 50, 51, 82 & 83 & 84

(keys), 76, 90

Notocorethrella 110

notoscriptus (Aedes, Finlaya, Culex,

Ochlerotatus) 5, 6, 50, 68, 80, 82

& 83 & 84 (keys), 103, 109, 113,

160 (map), 163, 165-168, 170, 172,

173, 176-178

novaezealandiae (Corethrella, Noto-

corethrella) 3, 7, 110, 111, 128

(figs.), 162 (map)

obscura (Dixina) 31

Ochlerotatus 50, 82 & 83 & 84 (keys),

Opifex 50, 51 & 52 (keys), 76, 80,

82 & 83 & 84 (keys), 93

otagensis (Nothodixa, Dixa) 3, 20, 21

(keys), 26, 28, 127 (figs.), 146

(map)

pacificus (Culex) 53, 54

Paradixa 14, 15 & 16 (keys), 29

Paradixini 15 & 16 (keys), 29

perkinsi section 81

pervigilans (Culex) 6, 7, 41, 50, 53,

54, 55 & 56 (keys), 64, 65, 66,

67, 69, 70, 81, 134 (fig.), 150 &

151 (maps), 163-178

pervigilans complex 53, 54, 69

perturbans (Coquillettidia, Culex) 74

philpotti (Nothodixa, Dixa) 3, 20, 21

(key), 24, 25, 41, 145 (map)

Phormium 67 —

pipiens (Culex) 53

pipiens complex 53, 54, 70

pipiens group 53

Potamogeton 175

quinquefasciatus (Culex, Culex pipiens)

5, 6, 10, 41, 50, 53, 54 & 55 & 56

(keys), 58, 60, 65, 68, 105, 153

(map), 167, 168, 170, 175, 176,

178

Rachionotomyia 108

*Ramcia 110

rotoruae (Culex) 50, 54 & 55 (keys),

98, 60, 64, 66, 67, 69, 133 (figs. ),

152 (map), 166, 175

Sabethini 106

Scirpus 167, 168, 169

septentrionalis (Nothodixa, Dixa) 3,

20, 21 & 22 (keys) 24, 27, 37, 126

& 127 (figs.) 146, (map), 167, 173

simplex (Nothopanax) 170

solomonis (Dixina) 10

Sphagnum 168

subalbirostris (Aedes, Ochlerotatus)

5, 81, 82 & 83 & 84 (keys), 100,

142 & 143 (figs.), 159 (map), 178

tamsi (Culex) 53

*tasmaniensis (Culex) 94

tasmaniensis (Paradixa) 32

tasmaniensis (Tripteroides, Rachiono-

tomyia) 108

tasmaniensis group 108

tenuipalpis (Coquillettidia, Austro-

mansonia, Taeniorhynchus, Man-

sonia) 3, 74, 75 (keys), 76, 78, 79,

136 & 137 (figs.), 155 (map), 167,

169, 177, 178

*Theobaldia 71

tonnoiri (Culiseta, Climacura, Theo-

baldia, Theobaldinella) 71, 154

(map), 171

tonnoiri (Paradixa) 15, 32, 33, 34

(keys), 36, 37, 41, 44, 127 & 131

(figs.), 149 (map), 170, 171, 175,

178

torrentium (Culex) 53, 54

trifilatus (Culex) 53

trifilatus subgroup 53, 54

Tripteroides 108

Typha 167, 175, 176, 177

unipunctata (Paradixa) 32

vagans (Culex) 53

vexans (Aedes) 113

vigilax (Aedes, Ochlerotatus) 102

vigilax section 81, 99

Contributions

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Volume 3, Number 2, 1968

MOSQUITO STUDIES (Diptera, Culicidae)

VI. A prodrome of the genus Orthopodomyia.

By Thomas J. Zavortink

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MOSQUITO STUDIES (Diptera, Culicidae)

Vill. A prodrome of the genus Orthopodomyia.

By Thomas J. Zavortink

oe ‘

aa aes

mee a

Nea ee

eek ea NS

MOSQUITO STUDIES (Diptera, Culicidae)

VIII. A PRODROME OF THE GENUS ORTHOPODOMYIA!

Thomas J. Zavortink2

CONTENTS

INTRODUCTION . Dea ett tw Mba i ial ea a ee eng hy enna en 2

et laee nest ii

MORPHOLOGY. .. . She ae Ose Nana, OPE er Ne aber had 6

BIOLOGY AND ECOLOGY . Se ae PAR ERD Er ey 8

DISEASE RELATIONSHIPS AND ECONOMIC IMPORTANCE Pg Be eye 9

Oe i ee eet Co a eet crea tty fig eee omens an

SYSTEMATICS. . gi let Rs oe ia Cee peng gear pete Pp Py SANG He toylca ge For

TAXONOMIC TREATMENT he i sidue hat Min Sif wea pet er he ee een gen et a

Grenue’Orthopodoms yids oa a ee ey pia ea lg ea Oa Ne

DANCTORUIA DOCTOR Cine caiecy oie ee OR OE eee

Siete re Cro ee ee Re re Sok Ls Sah cae

‘Signifera Subgroup . . . POI BE Sta gs AIRS A eae ats

1. Orthopodomyia waverleyi . POR ONE Sis Ua A eae ae

2. Orthopodomyia Sisnvera ieee oR ey ge

Ds Uy a ee gg ae

Pulchripalpis Subgroup |

4. Orthopodomyia PAECOE TADS it) APPS nahh BN Seite Ee

Kummi Subgroup. . . iis Wile ped aa att ee el cy |,

0. Orthopodomyia species ‘2 ie ie ane tpn hy he a he att

0. Orin pocon vin Buln aa a ee ae

Albicosta Group

4a, RED OOOINVIE BUCO ee ah. ca ee ae, Oe

Thomasina Section . . Sg te i. eae me ae

8. Orthopodomyia fascipes. Beg A aba ee ae

9. OTINGPGGDIMN Via BANIDALOL 50 lam a oe ee Oe

Pont pioatt it from project ''Mosquitoes of Middle America’ supported by

U.S. Public Health Service Research Grant AI-04379 and U.S. Army Medical

Research and Development Command Research Contract DA-49-193-MD-2478.

Based on PhD dissertation submitted to the University of California, Los Ange-

les and supported in part by U.S. Public Health Service Training Grant TI-AI-

132 and National Science Foundation Grant GB 3871.

y) :

Department of Zoology, University of California, Los Angeles, Califor-

nia 90024.

Z Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

Folicolae Section

10. Orthopodomyia DEY AU ECs peepee Le a a ce Be

Orthopodomyia Section. |... ee Se ge ae

Vernoni Group . . CeCe ee

il: Orthopodomyia Vernonia oie. ae 8G

22. OPO p0GO TIA TINO eo er ee 8

228 OTe op0comyia Sebel 6... a ee es 100

Nkolbissonensis Group

13. Orthopodomyia nkolbissonensis ......... . . 100

14. Orthopodomyia species a eo Ee

Arboricollis Group ;

1a. eee ALDOPICOLUS 6 oe 25 es

Albipes Group ...: . Co ae a ee

Flavicosta Subs roan. a ga as ae

16. Orthopodomyia flavicosta . A ear Oe

bis Orvibomogonivis avithorax 6. ck a ERD

“416. Orihopodomyia madrensis. . 3... 20... ss. « 122

19. Orthopodomyia Ssiamensis, n.Sp.< 2 os 220

Wilsoni Subgroup ,

2O. Orinonodoiayia wiisont <5 7 oes 2 ee

Albipes Subgroup |

aly Oi ooucoiyia dipe a see a eee ee Tee

Anopheloides Subgroup .. . a ee

22. Orthopodomyia papuensis, n. Sp. gop res aoe a

_/ 23. Orthopodomyia andamanensis......... 14!

24. Orthopodomyia anopheloides. ......... 146

: 3 ede Orthopodomyia meeresori. oS ee. ck TBA

Pe PERENCES fe Gi pole oe Fe ee ee ee ES

TUR | ee Pe ee a

INTRODUCTION

The mosquito genus Orthopodomyia is one of the poorer known genera in

the Culicinae (in the sense of Belkin, 1962). This is primarily because the

adults, with very few exceptions, are not attracted to and do not bite man;

consequently, the species are of no direct medical or economic importance and

have not received study by culicidologists. Another factor contributing to our

ignorance of the group is that all species breed in natural plant containers.

These habitats are often overlooked by mosquito collectors; thus, the imma-

ture stages of Orthopodomyia are seldom encountered unless specifically sought

and most species are poorly represented in museums.

In the past few years, however, a considerable number of specimens of

some species of Orthopodomyia have been collected in connection with mosqui-

to projects in Malaya, Thailand and Central America. As this additional ma-

terial, particularly of the immature stages, has become available, it has be-

come increasingly evident that the classification now existing for the group,

based largely on features of adult ornamentation, does not properly reflect the

Zavortink: Genus Orthopodomyia 3

relationships among the species.

The purpose, then, of the present study is to revise this genus from the

world standpoint on the basis of data from all known stages in the life cycle.

To achieve this end, species have been delimited on the basis of morphological,

ecological and distributional data and arranged into a classification which, it is

hoped, is consistent with the evolution of the group. Descriptions and illustra-

tions of all known stages of all species and descriptions of all higher taxa are

provided, keys to these taxa are included, data on the distribution and bionom-

ics of each taxon are Summarized, and speculations on the affinities of the var-

ious taxa are made.

Material for this study was first sorted to geographical origin and larval

habitat and then tentatively identified with existing keys or descriptions. I have

been fortunate in having either paratypic or topotypic material of a large num-

ber of species available for examination. My general procedure for any par-

ticular species was to first study this paratypic or topotypic material in consid-

erable detail. Then, when they existed and specimens were available, species

sympatric with the one being studied at or near its type locality were examined.

This process, essentially a reversion to that of the local naturalist dealing with

nondimensional species, was necessary to obtain an appreciation of the funda-

mental morphological differences between undisputed, sympatric species. Only

then were additional specimens, tentatively identified as being the species in

question, compared with the paratypic or topotypic material and a decision

made on the basis of morphological, ecological and geographical data as to

whether or not these individuals came from conspecific populations. This en-

tire process, when performed for each species, becomes somewhat self-cor-

recting and leads to a general refinement of one's concepts of the species in-

volved.

After species were delimited and diagnosed, they were grouped into higher

taxa of various levels on the basis of morphological similarity. Many of the

groupings are tentative only; until all the stages in the life cycle are known for

most species it will not be possible to construct a classification accurately re-

flecting the various affinities within the group.

I have examined specimens representing all previously described nominal

species except geberti, mcgregori and nkolbissonensis. Material contained in

the following institutions or private collections was studied: Bernice P. Bishop

Museum; British Museum (Natural History); California Academy of Sciences;

Canadian National Collection; Centre Scientifique et Technique, Bondy, France;

Cornell University; Fred Harmston; Instituto Oswaldo Cruz; Instituto de Salu-

bridad y Enfermedadas Tropicales, Mexico City, Mexico; Liverpool School of

Tropical Medicine; Antonio Martinez; Museo Firenze; South East Asian Mos-

quito Project, Washington, D.C.; United States National Museum; Universidad

de Sao Paulo; University of California at Davis; University of California at Los

Angeles; University of Utah.

Six of these collections deserve special mention because they have been

particularly vital to this study. The collections of the British Museum (Natu-

ral History) and United States National Museum have been most important be-

cause of their broad representation of species and accumulation of lectotypes

and holotypes. The excellent collection of William W. Macdonald has been in-

strumental in the treatment of the Oriental species because of its high quality

and great number of adults with associated larval and pupal skins. Important

also in the study of the Oriental species was the collection amassed by the South

East Asian Mosquito Project; this is the largest collection from that area. The

collection at UCLA has been invaluable, particularly in the study of the Neo-

4 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

tropical species. The bulk of this material has been collected for the project

on the Mosquitoes of Middle America. This collection is outstanding in its rep-

resentation of topotypic material, large series of specimens and number of in-

dividually reared adults. The last collection to be mentioned is my own. This

collection was important not because of itS representation of species but be-

cause I acquired an appreciation of the extent of intra- and inter-population var-

iation from it.

I wish to thank Dr. John N. Belkin for directing this study. Special thanks

are extended to Drs. P.F. Mattingly and J.A. Reid at the British Museum (Nat-

ural History) and A. Stone and B. de Meillon at the United States National Mu-

seum for the courtesies extended to me when I visited those institutions. Iam

also grateful to the individuals and institutions which loaned material. I am in-

debted to Sylvia Barr, Sally Dieckmann and Joyce Roberts for preparation of

the final plates and to Sheila Bernstein for typing the copy for reproduction.

HISTORY

The history of the classification of the genus Orthopodomyia is both short

and simple. The first species of Orthopodomyia was described as Culex pul-

chripalpis by Rondani in 1872 from material collected in Italy. In 1896 a sec-

ond species, the North American signifera, was described in the genus Culex

by Coquillett and in 1903 a third species, from India, was named Mansonia

anopheloides by Giles. During the next year 2 species were described and gen-

era based on them, Theobald (1904:236-237) creating the genus Orthopodomyia

for Leicester's Malayan species albipes and Lutz (1904) naming Bancroftia al-

bicosta from Brazil. Between 1905 and 1911, 9 additional species or varieties

of Orthopodomyia were described in the genera Orthopodomyia, Culex, Man-

sonia and Kerteszia and 3 additional generic names were published, Dyar (1905:

46) naming Pneumaculex for signifera, Theobald (1909:297) proposing Newstead-

ina for the Mauritian arboricollis, and Newstead and Carter (1911:553-554)

creating Thomasina for the South American longipalpis.

The classification of the family Culicidae was in a chaotic state during the

early twentieth century. Species and genera were being described at such a

rapid rate by workers throughout the world that many taxa were inevitably giv-

en more than one name. Additional confusion occurred because descriptions

were often so inadequate that many misidentifications and incorrect usages of

both specific and generic names took place. The greatest source of confusion,

however, was that many workers, particularly Theobald, created and diagnosed

numerous genera primarily on the basis of adult palpal length and scale mor-

phology. As a consequence of all these factors, even closely related species

were often not recognized as such and were placed in different genera. As an

example, in Theobald's (1910) A Monograph of the Culicidae, which sum-

marized the classification of the family, the 10 included nominal species of

Orthopodomyia were placed in the following genera: Orthopodomyia, Bancroftia,

Pneumaculex, Newsteadina, Mansonia, and Grabhamia.

Even before the initial period of proliferation of generic names was over,

a great reorganization and clarification of the classification of the Culicidae

started taking place. Genera were based on more fundamental morphological

features of adults in conjunction with characteristics of the larvae and male

genitalia. Fewer, but more natural, genera resulted.

In the case of Orthopodomyia, 4 of the 5 generic names proposed were re-

duced to synonymy and those species incorrectly placed in other genera were ~

Zavortink: Genus Orthopodomyia 9)

properly realigned. The initial steps toward this clarification were taken by

Dyar and Knab; first (1906:184) they reduced Pneumaculex to synonymy with

Mansonia and later (1910:264) they removed the orthopodomyias from Mansonia

and placed them in Bancroftia. The next step was taken by Edwards (1913: 239)

when he concluded that the New World Bancroftia was congeneric with the Ori-

ental Orthopodomyia. Howard, Dyar and Knab (1917:878) accepted Edwards'

concept of Orthopodomyia and in addition established that longipalpis was a syn-

onym of fascipes, thus reducing Thomasina to synonymy. Finally, Newsteadina

was synonymized with Orthopodomyia by Edwards (1920:135) when he examined

the type specimens of arboricollis.

Since 1911, 19 additional species or varieties of Orthopodomyia have been

described. All but two of these were described by people concerned only with

the mosquitoes of a restricted region (i.e., nondimensional species).

_ Although the species of Orthopodomyia have never before been the subject

of a thorough revision from the world standpoint, many species have been stud-

ied at one time or another in connection with regional faunal works. The most

important of these faunal works, which have included several species, are those

of Howard, Dyar and Knab (1917), treating the Central and North American ar-

ea; Dyar (1928), covering the entire New World; Barraud (1934); on the fauna

of India; Lane (1953), treating the Neotropics; and Carpenter and LaCasse

(1955), for America north of Mexico.

While these faunal works and many other studies dealing with 1 or 2 species

of Orthopodomyia have added to the knowledge of the species involved, they have

not materially contributed to an understanding of the relationships between the

species. There have been, however, three studies which have contributed to

the classification of the genus.

The first study is Edwards' classic treatise on the entire family Culicidae

(1932) in which one finds a brief treatment of Orthopodomyia from the entire

world. Solely on the basis of adult ornamentation he arranged the species into

2 groups, Group A (Orthopodomyia) and Group B (Bancroftia), and cataloged

the species in each. Except for the shifting of one species, phyllozoa, from

Group A to Group B (Knight and Mattingly, 1950), this classification of Edwards

has been accepted to the present time. The second study, by Edwards (1939),

slightly revised Group B (Bancroftia) and provided, by means of a key to the

adults of the included species, a classification of the group. The third study,

by Knight and Mattingly (1950), was of the anopheloides subgroup. This sub-

group, created by the authors on the basis of adult ornamentation, was a divi-

sion of Edwards' Group A (Orthopodomyia) which contained the Southeast Asian

species. Other subgroups proposed but not revised were the arboricollis and

fascipes. This study resulted in a better understanding of the relationships

within Edwards' Group A and among the species in the anopheloides subgroup.

A varying number of the nominal species proposed in Orthopodomyia have

been reduced to synonymy when the species have been studied in connection with

the larger faunal works. The synonymies established or expressed in Knight

and Mattingly (1950) and Lane (1953) are currently accepted in Stone, Knight and

Starcke's world catalog (1959).

In summary, the situation at the start of the present study is that 33 spe-

cific names are available in Orthopodomyia and 26 of these are considered to

represent valid species or subspecies. On the basis of adult ornamentation the

species are arranged into 2 groups, Group A (Orthopodomyia) and Group B

(Bancroftia), and the species in the Orthopodomyia group are placed in 3 sub-

groups.

6 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

MORPHOLOGY

This section is not an exhaustive description of a representative species of

Orthopodomyia, but is instead a discussion of some characteristics within the

genus Orthopodomyia which have been found particularly useful or too variable

for taxonomic purposes. In this discussion, and in the descriptions of the

groups and species, the beh iamiees 2! and abbreviations used are those employed

by Belkin (1962:547-563).

The adults of Orthopodomyia are all very similar in structural details.

The head morphology, except for small differences in the palpal length of each

sex and the number of palpal segments of the female, is similar in all species

except those in the ThomaSina section. Even in the Thomasina section the dif-

ferences, restricted to males, are not extensive; the torus is reduced in Size,

the flagellum has fewer and shorter bristles and the proboscis is stout and

strongly sclerotized. The palpus of the female is 4-segmented in all species

except for some in the Albipes and Signifera groups which have a 5-segmented

palpus. The external anatomy of the thorax is nearly the same in all species.

Group characters are found in the number of pra and stp bristles. Species in

the Albipes group are the only ones without a single strong upper stp bristle

_ and the species in the Signifera group have an increased number of pra bristles.

The species in the Signifera group are the only Orthopodomyia to have the first

hindtarsal segment shorter than the hindtibia.

Except for the morphological differences just discussed, adults of the spe-

cies and groups of Orthopodomyia differ only in ornamentation. Particularly

valuable group characters are found on the thorax and wing. While the differ-

ences between the groups are, of course, constant, it is often difficult to find

reliable specific differences. Ornamentation of the proboscis, palpus, legs,

wing and abdomen is extremely variable. Part of this variation, at least in the

Signifera group, is correlated with larval nutrition and some appears to be en- |

vironmentally induced.

The male genitalia of all Orthopodomyia are also very similir’ Specific

and group differences are largely restricted to the shape, sclerotization and

dentition of the aedeagus. Only the male genitalia of the Thomasina sectionare

"conspicuously" different; in this group the basal mesal lobe has 2 or 3 stout,

short, blunt and flattened bristles and the eighth tergite does not have a lobe on

the posterior margin. The Nkolbissonensis group may differ in development of

the bristles on the ninth tergite. Other group characters are found in the spini-

form of the clasper; the Thomasina section and the Vernoni and Signifera groups

have a lobed spiniform and the Albicosta group has 2 simple spiniforms. The

shape of the lobe on the eighth tergite is quite variable and cannot be used ex-

tensively as a taxonomic character. The number of bristles on the basal mesal

lobe and the ninth tergite is not characteristic of species or, eee as noted

above, groups.

The pupae and larvae of Orthopodomyia each conform to one morphological ©

plan. There are no differences in the pattern of chaetotaxy between species or

groups. For the most part, the taxonomic characters are restricted to the

length and number of branches of the hairs. It is difficult or impossible to find

specific differences in some cases because of the variability of both these fea-

tures. This is best illustrated by the Signifera group, in which the majority of

the populations of each allopatric species has a characteristic development of

certain hairs in the continuum from weakly developed to strongly developed,

but in which there are enough individuals and populations within each species

Zavortink: Genus Orthopodomyia 1

with these hairs more or less strongly developed than normal to prohibit diag-

_ nosis of the species on the basis of chaetotaxy. Many species of Orthopodomyia

have "hairy’’ forms in which the development of many or most body hairs is

strikingly changed. Normally short single hairs may become extremely long

and multiple and short multiple hairs may become long and fewer branched. In

at least 1 species, arboricollis, such changes in the chaetotaxy of the larva are

accompanied by changes in the length and curvature of the antenna.

Except for differences in chaetotaxy, the pupa has few characters of taxo-

nomic importance. The development of hairs 1-7-C and 9-VII, VIII, and the

position of hairs 2-VI, VII have been found to be group characters. Pupae of

the species in the Albipes group may differ in paddle and trumpet shape; while

these differences are not striking, they do appear to be constant.

The evolution of the head morphology of Orthopodomyia larvae has been

conservative. Only phyllozoa has a unique head capsule shape and only the spe-

cies alba, arboricollis and flavithorax have conspicuous differences in the

length and number.of branches of some head hairs. By contrast, specific and

group characters can usually be found on the terminal segments of the larva.

The size, shape, number and arrangement of the comb scales are good group

characters and the shape of the comb scales is a specific character in the Al-

bipes group. The number of comb scales is too variable to be useful at the

specific level; in several instances, both extremes for the range in the number

of comb scales in a particular species have been encountered in 1 individual.

The siphon and anal segment often provide both group and specific characters.

Particularly important are the length of the siphon, the development of hair

1-S, the number of hairs in the ventral brush and the length of the anal gills.

Additional group and specific differences are found in the chaetotaxy of the tho-

rax and abdomen. Important group characters are the development of thoracic

hairs 1, 4-M and 1-T, and the position of abdominal hairs 2-VI, VII.

Several features of the larva are variable enough to deserve mention.

First is the development of the sclerotized plates on segments VI, VII and VIII

of the fourth instar larva. It has been shown by Tate (1932:115), Reeves (1941:

70-71) and Chapman (1965:436) that these plates are added during the growth of

the last instar larva. This fact was also known by Howard, Dyar and Knab

(1917:893). As a consequence, the plates can be expected to vary in specimens

which may have been killed at different ages. Specimens at the same physiolog-

ical age (pupation) also vary in plate development. Chapman found 62 9o of the

larval skins he examined with plates on segments VII and VIII and 38 9% with

plates on segments VI, VII and VIII. Examination of skins from larvae collec-

ted at 2 different times in the same treehole has shown that winter-collected

diapausing larvae usually have a more extensive development of plates than

summer-collected non-diapausing larvae. The variability of the extent of these

sclerotized plates is strikingly shown by occasional specimens which have large

plates on the thorax. Two other areas of sclerotization are involved in age var-

iation. The first is the presence and size of the ring basad of the anal saddle 3

and the second is the presence or absence of an acus. Young fourth instar lar-

vae do not have a ring basad of the saddle, but as the larvae grow a sclerotized

band forms on each side and those from opposite sides may sometimes join to

form an incomplete ring. Young fourth instar larvae often have a detached ac-

us which, as the sclerotization of the base of the siphon increases with age,

first becomes attached and is finally completely incorporated into the base of

the siphon. The pigmentation of the larva is also quite variable. Some species

have strongly developed red or purple epidermal pigments. These pigments

8 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

mask the color of the fat body and the larvae do not conspicuously change color

with age or diet. Other species either do not have or have only weakly devel-

oped epidermal pigmentation, and in these the color of the fat body becomes

conspicuous. These larvae are usually pink when young but become bluish as

the time of pupation approaches. They may also become yellowish on labora-

tory diets. In addition to these biological variations, some mounting media.-re-

move or change the color of the epidermal pigment.

BIOLOGY AND ECOLOGY

The following discussion is based on observations made in the field in Ari-

zona, southern California, New Mexico, Ohio, England and Guatemala, and

those made in the laboratory. The species studied were alba, kummi, pulchri-

palpis, signifera and waverleyi.

Orthopodomyias are usually sought and collected as larvae, which, in the

areas Studied, are found intreeholes. Individuals of all instars may be found

in any month of the year. During the winter, first, second, third and small

fourth instar larvae become quiescent, but mature fourth instar larvae go into

diapause. The larvae are not found in all treeholes (see below), but they may

be present in large numbers in the proper type of hole.

The type of hole in which the immature stages are found depends upon the.

local climate. In cismontane southern California, females are active from

May until October or early November. Since this flight period coincides with

the dry season, only very few treeholes, the ones with permanent water, are

available for oviposition. The permanent treeholes are of two types. Some are

just particularly deep holes with small openings in oak (Quercus) trees; these

holes are filled during the rainy season and although the water level drops con-

siderably during the dry season they never quite dry out. The other type of

permanent treehole is found in some species of cottonwood (Populus) and wil-

low (Salix) trees. These holes are filled by fluids from the tree and the water

level is independent of precipitation. Temporary treeholes, filled only during

and for a short time after the rainy season, contain water at a time when there

are no ovipositing females and as a consequence never contain Orthopodomyia

larvae. The situation is somewhat different in the mountains in southeastern

Arizona because there are 2 rainy seasons, a lesser one from December to

February, and a greater one in July and August. As a consequence of being

filled twice a year, a greater proportion of the oak treeholes contain water per-

manently and Orthopodomyia larvae can be found in them at any time. In addi-

tion, Since the rainy season in July and August falls within the period of adult

activity, many temporary treeholes will contain Orthopodomyia larvae at that

time. The immature stages are, of course, also found in the permanent cot-

tonwood and willow treeholes.

Various comments in the literature lead me to believe that most, if not all,

species of treehole breeding Orthopodomyia are found more frequently in per-

manent than temporary treeholes. Particularly important as Orthopodomyia

breeding sites may be those tree species in which the water in the hole is sup-

plied by the tree and is independent of precipitation. In addition to the cotton-

woods and willows mentioned above, this phenomenon is also found in horse-

chestnut (Aesculus hippocastanum), elm (Ulmus), and beefwood (Casuarina)

trees. The fluid in these holes kept filled by the tree is usually lighter in color

(sometimes straw-colored or even colorless) and more alkaline than that of

Zavortink: Genus Orthopodomyia 9

other treeholes. Permanent treeholes seem also to be the preferred breeding

places of treehole Toxorhynchites and Anopheles, and as a consequence these

genera are frequently found with Orthopodomyia.

Orthopodomyia larvae are apparently filter feeders, ingesting microorgan-

isms and small particles which they have strained from the water with their

mouthbrushes or plumose body hairs. They are rarely seen gnawing at large

objects, browsing over the surface of submerged objects, or feeding from the

surface film.

Larvae can easily be reared in the laboratory in diluted treehole water to

which yeast is added if the medium is kept alkaline (as is the water in treeholes)

by addition of sodium hydroxide. Despite being crude, this method is very suc-

cessful, with the larvae growing rapidly and producing large, robust adults.

The pupal stage of Orthopodomyia lasts from 5 to 8 days.

- Very little is known about the adults of Orthopodomyia. All specimens en-

countered in the field were resting above the water level on the inside of tree-

holes. They were quite wary and often flew away at the approach of an observ-

er. While most usually settled on the bark near the treehole, others flew across

sunlit areas to nearby trees. All specimens caught were freshly emerged.

Laboratory observations indicate that the adults are active only after dark-

ness. Males seldom live more than 2 weeks and females fed on sugars live 4

to 6 weeks. Females of alba, signifera and kummi readily took blood meals

from a canary (Serinus canaria), brown towhee (Pipilo fuscus), and a house

sparrow (Passer domesticus), but would not feed on a pigeon (Columba livia).

None could be induced to feed on a human arm. Gravid females oviposited dur-

ing darkness on moist filter paper lining containers partially filled with filtered

treehole water. Eggs were laid singly or in small groups; those of signifera

and kummi were deposited within 3 or 4 mm of the water meniscus, but those

of alba extended to 3 or 4 cm above the meniscus.

Of several thousand eggs obtained, only about 60 (of signifera) hatched. All

others checked showed no development and were apparently never fertilized.

Those which hatched did so immediately after embryonic development and did

not go into diapause. :

DISEASE RELATIONSHIPS AND ECONOMIC IMPORTANCE

Only 2 species of Orthopodomyia, the Oriental albipes and andamanensis,

are known to take human blood. Since neither has been implicated as a vector

of human disease or bites man frequently enough to be considered a pest, they,

and all other species of Orthopodomyia, are usually dismissed as not having

medical or economic importance. However, Orthopodomyia may be of consid-

erable indirect medical importance. Three species, the Nearctic alba, kummi

and signifera, are known to attack birds and are possibly important in the

maintenance of arboviruses in sylvan environments or in the transmission of

them to domestic fowl. Vargas (1960:339, 340) reported isolating the virus of

Eastern Equine Encephalitis from signifera in Mexico. Chamberlain, Sikes,

Nelsen and Sudia (1954:283) gave signifera ''good" and "excellent" vector po- |

tential ratings for Eastern and Western Equine Encephalitides respectively.

An appreciation of the role played by Orthopodomyia as vectors of arboviruses

cannot be acquired until the biology of a greater number of species is investi-

gated.

10 Contrib. Amer. Ent. Inst., vol. 3) no. 2, 1968

DISTRIBUTION

Fig. 1 |

Orthopodomyia has been enly sparsely collected and the geographical dis-

tributions of the species are very poorly known. The data which are available

are summarized here.

The genus is worldwide in distribution, but is largely restricted to tropical

and warm temperate areas. In addition, each of the 8 species groups in the

genus is entirely or largely restricted to one faunal region.

_ A large number of species, belonging to one diverse group, iS concen-

trated in the Oriental region. Some species of this group have spread as far

as Japan, New Guinea and northeastern Australia (Marks, 1962, in litt.).

While the Neotropical region has fewer species than the Oriental, it has

the greatest representation of distinct groups, with 4 being present. One of the

groups has spread northward into the Nearctic and Palearctic regions; this

represents the only known large scale invasion of colder areas by Orthopodo-

myia species.

Very little is known about the Orthopodomyia fauna of the Ethiopian region.

Muspratt (1955:156) was the first to indicate the presence of a species of Ortho-

podomyia in Africa, having found specimens in Kruger National Park, South

Africa. This species has not yet been described. In early 1967 a species was

described from Cameroon; it appears to represent a distinct group. How many

species or groups actually occur in Africa is entirely problematical. Accord-

ing to Grjebine (1966:29, 30, 32), the island of Madagascar has 7 species, only

3 of which have been described, belonging to one group and the island of Mayo-

te has one undescribed endemic species of the Same group. The island of Mau-

ritius, in the Indian Ocean east of Madagascar, has one endemic species of Or-

thopodomyia. This species represents a distinct group.

The breeding sites of Orthopodomyia could theoretically limit their geo-

graphical distribution. This does not appear to be the case, however, since

angiosperm trees, bamboos, bromeliads and heliconias hve more extensive

distributions than those species breeding in them.

Although the majority of species breed in treeholes, the genus is not re-

stricted to mesic, extensively forested areas. One species is often encoun-

tered in holes in cottonwood (Populus) and willow (Salix) trees along the water-

courses in the Mojave, Colorado and Sonoran Deserts in the western United

States. : |

SYSTEMATICS

Data from morphology, ecology and geography have been uSed in the inter-

pretation of the systematics within the genus. Primary emphasis has been

placed on the comparative morphology of the adults, male genitalia, pupae and

larvae. The most important criterion has been that each species be character-

ized by several correlated morphological characters. Except for the species

in the Signifera group, these correlated characters are found in all 3 stages

and the male genitalia; within the Signifera group ee are largely restricted

to the adult stage.

The basic ecological criterion which I have used is that the immature stages

of not more than one species in each compléx will be found occupying the same

habitat in any given area. This criterion has been extremely important because

Zavortink: Genus Orthopodomyia 11

the chaetotaxy of the immature stages of Orthopodomyia is often modified into

"hairy forms, ‘'' in which several correlated morphological characters are ex-

pressed in larvae, pupae, or both.

There are, however, 4 apparently bona fide instances in which the larvae

of species in the same complex do occupy the same habitat in the same area.

In 2 of the cases the larva of one of a pair of closely related sympatric species,

both of which breed in treeholes, is very extensively modified. The species

pairs involved are alba and signifera in North America and flavicosta and flavi-

thorax in India. I believe that these 2 cases, rather than being exceptions to

the ecological tenet, offer striking evidence of its validity; only in those cases

where extensive secondary modification of the larva (the feeding stage) of one

species has taken place are 2 species in the same complex enabled to coexist

indefinitely over a broad area. In the third case, the larvae of the species oc-

curring together are very similar. But, the species involved, kummi and Sig-

nifera, are basically allopatric and are known to occupy the same treeholes in

only one area. It is possible that it is the similarity of their larvae which has

prevented either of these species from penetrating the range of the other. In

the fourth case the species involved, anopheloides and andamanensis, have very

Similar larvae and yet are apparently sympatric over a broad area. Possible

explanations are that andamanensis is not a distinct species, but is only a gen-

etic segregate of anopheloides, or that the environment is diverse enough in

time or space that neither species has eliminated the other.

Data from distribution, that is, allopatry or sympatry, have been used,

but no rigid rules have been followed in deciding what rank to give morpholog-

ically distinct allopatric or sympatric populations. In general, if allopatric

populations are characterized by the same amount of morphological difference

as the populations of sympatric species, then these allopatric populations have

been considered to represent distinct species. If, however, the differences

are of lesser magnitude, the populations are usually considered to be conspe-

cific. I have not recognized any subspecies because in most instances so few

Specimens are available that one does not know whether the differences ob-

served are truly characteristic of populations of an area or are individual or

environmentally induced variation. Sympatry of entities morphologically dis-

tinct by several correlated characters has been interpreted as indicating spe-

cific rank for the taxa involved.

The classification of Orthopodomyia which follows is based on features of

adults, male genitalia, pupae and larvae. A classification based on any one of

these stages would differ from that based on any other; I do not necessarily

mean to indicate that these classifications would be conflicting, only that they

would be different.

: There are, however, several instances in Orthopodomyia in which the

classification based on adult ornamentation is at variance with that based on

the immature stages. Depending upon the situation, I have in Some cases de-

cided that the relationships shown by the immature stages were the true ones

but in other cases have given precedence to adult characters.

In general, I have found the pupal stage to be the most reliable both for

showing specific differences and affinities. It is superior to the larva for 2 |

reasons. First, the pupal chaetotaxy is less often and less extensively modi-

fied into hairy forms than is that of the larva (except in the Anopheloides sub-

group); and second, since the pupa is not a feeding stage, it is not subject

to selection pressures reducing competition for food, and consequently does

not show the extensive modifications of morphology that some larvae do (e.¢g.,

alba and flavithorax).

12 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

In this study 24 species in 8 distinct groups are recognized in the genus

Orthopodomyia. The groups, with their distribution and included species, are

as follows: (1) the Oriental Albipes group with the 9 species, flavicosta, flavi-

thorax, madrensis, siamensis, wilsoni, albipes, papuensis, andamanensis and

anopheloides; vides; (2) t the Holarctic Signifera eroup with the 6 species waverleyi,

signifera, alba, pulchripalpis, species 5 and kummi; (3) the Neotropical Thom-

asina section with the 2 species fascipes and sampaioi; (4) the Madagascan

Vernoni group with the 2 species vernoni and milloti; (5) the African Nkolbis-

sonensis group with the 2 species nkolbissonensis and species 14; (6) the mono-

typic South American Albicosta group; (7) the monotypic Mauritian Arboricollis

group; and (8) the Middle American Folicolae section with the single species

phyllozoa. :

At the present time the affinities of these 8 groups are not obvious; they

may become evident, however, when the immature stages, which are at least

partially unknown for half the groups, are described. The groups have been

placed provisionally into 4 sections, as follows: (1) the Orthopodomyia section

made up of the Albipes, Vernoni, Nkolbissonensis and Arboricollis groups; (2)

the Bancroftia section containing the Signifera and Albicosta groups; (3) the

Thomasina section; and (4) the Folicolae section.

_ The primary mechanism of speciation in Orthopodomyia appears to be geo-

graphical isolation. The species in the Signifera group, the Flavicosta sub-

group and the Thomasina section are entirely or largely allopatric. The species

in the Vernoni and Nkolbissonensis groups may also be allopatric. There is no

evidence of sympatric ecological speciation in Orthopodomyia; closely related

species are either allopatric or sympatric in the same habitat. In those cases

where 2 closely related species are sympatric in the same habitat, I have as-

sumed that the species were originally geographically isolated. With the possi-

ble exception of the Albipes group, complete hybridization does not appear to

have been a mode of speciation in Orthopodomyia. The Anopheloides subgroup

of the Albipes group may have arisen through hybridization between species in

the Flavicosta and Albipes subgroups. Such an interpretation would not only

explain some aspects of the morphology of the immature stages of the Anoph-

eloides subgroup, but would also explain a portion of the discordance between

adult and larval classifications in the Albipes group.

Although complete hybridization does not appear to have been an important

mode of speciation, introgressive hybridization has evidently played a major

role in the evolution of the genus. One of the most remarkable phenomena to |

be discovered in this study is that a species which contacts or almost contacts

- another often has characteristics, particularly in the adult, of the latter spe-

cies. It appears that at least a portion of the cases where characters are dis-

tributed in this way can be explained only by introgressive hybridization, and

that phenomenon must accordingly be considered very important in the evolu-

tion of Orthopodomyia. Introgression can apparently take place between spe-

cies in the same group or between those of separate groups; it has possibly oc-

curred between the following species: (1) kummi and signifera, (2) kummi and

albicosta, (3) kummi and fascipes, (4) fascipes and phyllozoa, (5) pulchripalpis

and some species in the Anopheloides subgroup, (6) flavicosta and flavithorax,

and (7) between various species in the Albipes group.

There is one particular instance in which the evidence for introgression is

especially strong and I think it is worthwhile to elaborate this case here. The

South American Albicosta, Oriental Albipes and Holarctic Signifera groups are

involved. The Signifera group has several apomorph characters not found in

Zavortink: Genus Orthopodomyia : 13

the other 2 groups and since it has widespread allopatric species while the other

2 groups each have a species or subgroup with a relict distribution, it appears

to. be more recent than the Albipes and Albicosta groups on the basis of distri-

bution. It is important to stress these factors because the argument for intro-

gression is based on the fact that neither the Albicosta nor Albipes group could

have been derived from the Signifera group. The Signifera group is centered in

North America but has one species in Europe (pulchripalpis) and one in Central

America (kummi). Pulchripalpis has several features which are unique for the

group but which are found in the Oriental Albipes group and kummi has several

traits which are found elsewhere only in the South American Albicosta group.

While one could postulate the derivation of the Signifera group from either the

Albipes or the Albicosta group, the similarities between the Signifera group and

the remaining group may be due to introgression since the latter group cannot

be derived from the former. I have considered the similarities between kummi

and the Albicosta group to be due to relationship and those between pulchripal-

pis and the Albipes group to be due to introgression. It is possible, however,

that the similarities between kummi and the Albicosta group are also due to in-

trogression.

Hybridization between species must, of course, occur before introgression

can take place. Evidence of hybridization between 3 different pairs of species

has been seen during this study; the pairs involved are kummi and signifera,

flavicosta and flavithorax, and albipes and anopheloides.

I do not believe that the evidence for hybridization or introgression in any

way militates against considering the taxa involved as distinct species. Despite

gene exchange, these taxa appear to be separate lineages with their own roles

and tendencies and are accordingly given specific rank.

TAXONOMIC TREATMENT

The species have been aggregated into numerous groups at various levels |

to indicate their affinities, and both species and groups (but not the species

within each group) have been arranged so that the most generalized or primi-

tive are placed centrally and the most derived are placed at the ends.

Descriptions are extensive because the genus is poorly known and many

undescribed species or forms probably exist. Species descriptions are com-

posite, that is, based on many individuals. Group characters are not repeated

in the descriptions of included taxa.

Complete synonymy is provided for all species group and generic group

taxa. An extensive list of literature citations is provided for each species.

Those taxa for which the type specimen has been seen are marked by an aster-

isk.

The systematics section of each species or group contains restatements or

explanations of diagnostic characters, explanation of the treatment, discussion

of geographic or other variations, mention of hybrids, and speculations on evo-

lution and relationships.

The abbreviations used in the lists of specimens examined are those of

Belkin (1962:563-564) or Stone, Knight and Starcke (1959:327-329) except for

the following: Fred Harmston [HARM]; Antonio Martinez [MART]; University

of California at Davis [UCD].

14 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

Genus ORTHOPODOMYIA Theobald

1904. Orthopodomyia Theobald, 1904:236-237. TYPE SPECIES: *Orthopodo-

myia albipes Leicester, 1904, Malaya; monobasic.

1904. Bancroftia Lutz, 1904:6,1. TYPE SPECIES: Bancroftia albicosta Lutz,

1904, Brazil; monobasic. :

1905. Pneumaculex Dyar, 1905:46. TYPE SPECIES: *Culex signifer Coquillett,

1896, United States; original designation.

1909. Newsteadina Theobald, 1909:297. TYPE SPECIES: *Culex arboricollis

Charmoy, 1908, Mauritius; original designation.

1911. Thomasina Newstead and Carter, 1911:553-554. TYPE SPECIES: *Man- |

sonia longipalpis Newstead and Thomas, 1910, Brazil; original designa-

tion. .

FEMALES. Small, medium-sized or large mosquitoes, varied in appear-

ance, ornamented conspicuously with narrow lines of white scales or broader

lines of silver and golden scales on a dark integument (Bancroftia and Folicolae

sections respectively), or less conspicuously with a general mottling of pale

and dark scales (Orthopodomyia and Thomasina sections), readily distinguished

from all other mosquito genera by the unusual proportions of the tarsal seg-

ments on the fore- and midlegs, segment 1 longer than the remaining 4 com-

bined and segment 4 shorter than segment 5. Head: eyes narrowly separated

above antennae, partially contiguous (Signifera group, in part), or entirely con-

tiguous (albicosta); frontal tuft absent; frontal bristles 1 pair, strongly devel-

oped; orbital bristles rather few, lateral ones moderately developed, directed

cephalad, mesal ones strongly developed, directed over the bases of the frontal

bristles; vertex and occiput with few to numerous decumbent scales, all narrow

and curved or with some broad and flat (albicosta), and numerous to very nu-

merous erect scales; dorsolateral, lateral and ventral surfaces with broad flat

decumbent scales; suborbital bristles numerous, weakly developed; ornamen-

tation varied; clypeus small, an elongate angular lobe or triangular (albicosta,

Kummi subgroup), normally without bristles or scales; labium quite broad,

slightly to considerably swollen apically, with several subbasal bristles, shaft

bristles absent or weakly developed, ornamentation varied; labella scaleless;

palpus 0. 33 to 0.58 of proboscis, bare of scales ventrally, 4- or 5-segmented,

segment 4 moderately long to very short, segment 5 short or minute, segments

2 and 3 with well developed bristles, ornamentation varied; antennae longer

than proboscis, scape scaleless, torus enlarged, flagellum 13-segmented, all

segments subequal or first and last slightly longer, first sometimes somewhat

swollen (albicosta, phyllozoa), flagellar whorls consisting of relatively few

bristles arising from near base of each segment. Thorax: mesonotum moder-

ately arched; scutellum trilobed; postnotum bare; paratergite narrow to broad,

rarely with scales (flavicosta, flavithorax); apn lobes medium-sized, widely

separated; pra not separated by a definite suture from stp; meron large, its

upper edge well above base of hindcoxa; acrostichal, dorsocentral and lateral

prescutellar bristles numerous, very strongly developed, acrostichals ina

more or less double row from anterior promontory to cephalad of prescutellar

space, dorsocentrals and lateral prescutellars forming a continuous line from

anterior promontory to near scutellum, lateral prescutellars and sometimes

posterior dorsocentrals in a more or less double row; humeral, lateral pre-

scutal and posterior fossal bristles present, all strongly developed or some

weakly developed, sometimes numerous enough to enclose fossa or with pos-

Zavortink: Genus Orthopodomyia 15

terior fossal bristles absent (albicosta); fossal bristles present or absent; sup-

raalar bristles numerous, strongly developed, in 2 more or less distinct rows

mesad of paratergite, in a single broad row over wing base; parascutellar bris-

tles normally 1; prescutellar space bare except for fine bristles in center; lat-

eral scutellar lobes with bristles numerous, some strongly developed, midlobe

with bristles few, more or less divided into 2 groups, some strongly developed;

mesonotum with narrow curved decumbent scales, ornamentation varied; scu-

tellum with large and sometimes small decumbent scales; apn with numerous

weakly to strongly developed upper anterior bristles and usually with weakly to

strongly developed lower bristles; ppn usually with 2 or 3 strongly developed

posterior bristles and sometimes with 1 or 2 additional weakly developed ones,

normally without upper bristles; ppl bristles numerous, weakly to strongly de-

veloped; sp and psp bristles absent; pra bristles 0, 1, or 2-10 (Signifera group),

moderately to strongly developed; upper stp bristles 0, 1 or 2, moderately to

strongly developed, posterior stp bristles 0, 1, 2, or several, weakly to strong-

ly developed, lower stp bristles 0 or several, weakly to strongly developed, stp

always with at least posterior or lower bristles developed; mep with upper pos-

terior bristles few to numerous, moderately developed, anterior bristles rare-

ly present (pulchripalpis); pleuron with narrow curved and broad flat decumbent

scales, ornamentation varied. Legs: fore- and midfemora of greater diameter

than hindfemur; femora with inconspicuous to conspicuous erect scales ventral-

ly at apex; fore- and midtarsal segment I longer than the remaining 4 seg-

ments combined, segment 4 markedly shortened, shorter than segment 5; hind-

tarsal segment 1 longer or shorter than hindtibia; claws of all legs medium-

sized, subequal, simple, or hindclaws smaller; pulvilli not developed; fore-

coxa with numerous strongly developed bristles along outer anterior edge, mid-

coxa with nearly vertical row of strongly developed bristles on outer margin,

hindcoxa with strongly developed bristles along lower outer posterior edge, all

coxae with additional bristles associated with coxa-trochanter joint; trochanters

without bristles; femora and tibiae with well developed bristles; ornamentation

varied. Wing: wing membrane with distinct microtrichia; scales symmetrical —

or asymmetrical, broad or narrow, lying close against the veins or spreading;

ornamentation varied; vein Sc ending between level of base of vein R4+5 and

level of furcation in vein R9,3, base without bristles ventrally; remigial bris-

tles on base of vein R dorsally present or absent; veins R, and R445 without

basal spurs; cell Rog varied in length but usually longer than vein Rg+3; cross-

vein r-m far distad of crossvein m-cu; cell Mg varied in length but usually

longer than vein M measured from crossvein m-cu to its furcation; plical vein

strongly developed, with or without scales at base ventrally; vein 1A smoothly

curved or slightly sinuate except at apex where it turns sharply towards poste-

rior margin, apex far distad of furcation in vein Cu; costa concave between ends

of veins Cuz and Cug; fringe without light spots; alula with a few narrow to me-

dium-wide marginal scales; upper calypter with continuous marginal row of

bristles or scalelike bristles, lower calypter bare. Haltere: short, stubby,

knob and anterior edge of stem with small flat scales. Abdomen: bristles pres-

ent on tergites I-VIII and sternites II-VIJ; tergite I with middorsal scales and

with or without scattered scales; laterotergite with or without scales; tergites

II- VIII completely and densely scaled; sternite Imembranous, bare; sternites

II-VIII not densely scaled. Buccopharyngeal Armature: not developed.

FEMALE GENITALIA. Segment VIII not retractile, sternite longer than

tergite, with straight posterior margin; tergite IX distinct, narrow, extending

far ventrad, with or without dorsolateral setae; tergite X not developed as a

16 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

distinct entity; cercus large, extending beyond postgenital plate, strongly com-

pressed, scaleless, with apical setae; postgenital plate slightly longer than

broad, deeply notched apically, resulting lobes with strong setae; cowl strongly

sclerotized, fused to cephalic portion of postgenital plate; sigma weakly sclero-

tized; insula weakly sclerotized, connected to sigma, bare or with a pair of

minute setae; spermathecae 3, more or less spherical, strongly sclerotized,

1 slightly larger than other 2. | ,

MALES. Essentially as in females. Labium: not more strongly swollen

than in females or of much greater diameter throughout length than in females

and strongly sclerotized (Thomasina section). Palpus: 0.75 to 1.02 of probos-

cis, 5-segmented, segments 2 and 3 more or less ankylosed and forming about

0.67-0.75 of palpus, segment 5 short to very short, segments 4,5 and some-

times 3 with a few well developed bristles. Antenna: subequal to or longer than

proboscis, torus moderately (Thomasina section) to greatly enlarged, flagellum

with segment 1 slightly elongated, segments 12 and 13 greatly elongated, mod-

erately (Thomasina section) to strongly plumose, bristles on segments 1-12 very

dense, arising from or distad of middle of segment on segments 1-11, but near

base on segment 12, those on segment 13 sparse, arising near base of segment.

Legs: anterior fore- and midclaws large, biserrate, basal tooth large (Thom-

asina section) or too small to be seen on pinned specimens, posterior fore- and

midclaws medium-sized, simple or uniserrate with tooth too small to be seen

on pinned specimens (Bancroftia section), hindclaws small, subequal, simple.

Abdomen: bristles slightly stronger and more numerous, particularly on ster-

nites, than in females; sternite VIII densely scaled. |

MALE GENITALIA. Segment VIII: long and wide; tergite bearing a distinct

median lobe on posterior margin or posterior margin only slightly rounded

(Thomasina section); sternite unspecialized. Segment IX: medium-sized; ter-

gite poorly sclerotized, without lobes, with or without setae dorsolaterally;

sternite normally without bristles. Sidepiece: well developed, usually more or

less long conical or cylindrical; basal mesal lobe small, more or less distinct,

bearing specialized apical bristles, some of which may be shortened and flat-

tened (Thomasina section); no other lobes developed; mesal surface completely

sclerotized or partially membranous; mesal and/or tergomesal surfaces some-

times bearing specialized bristles distad of basal mesal lobe. Claspette: rep-

resented by basal mesal lobe of sidepiece. Clasper: simple, slender, arising

subapically on sidepiece; very fine setae present, more numerous distally.

Spiniform: usually 1, rarely 2 (albicosta), subapical on clasper, simple or with

apex flattened, broadened, partially rolled, and divided into regular or irregu-

lar lobes. Phallosome: simple; aedeagus large, strongly or weakly sclero-

tized, varied inform, with or without apical, basal, tergal or sternal teeth,

Spines or processes; ventral paramere strongly developed, articulating with

basal piece and aedeagus; dorsal paramere not developed; basal piece strong-

ly developed, not joined to sidepiece, but articulating with it, with ventral para-

mere and with basolateral sclerotization of proctiger. Proctiger: strongly de-

veloped; basolateral sclerotization prominent, articulating with tergite IX and

basal piece; paraproct sclerotization prominent, with 1 to several apical teeth;

tergal surface not sclerotized; cercal sclerites absent, cercal setae few, very

fine, distinct. |

PUPAE. Cephalothorax: middorsal ridge strong; all hairs present, vari-

ously developed; hair 6-C always smaller than 7-C; 6, 7-C close together, hair

6 usually cephalad of hair 7, rarely caudad of it (flavicosta, flavithorax); 8, 9-C

relatively close together and always far caudad of trumpet. Trumpet: not placed

Zavortink: Genus Orthopodomyia 17

on distinct tubercle; widely spaced, much nearer wing base than middorsal line;

moderate in length; tracheoid absent, entirely reticulate; distal portion gradu-

ally widened from base in most species; meatus without slit; pinna small to

large. Metanotum: fourth pair of hairs rarely developed (flavithorax). Abdo-

men: all normal hairs except 10,11-I present; hair 1-I strongly developed and

strongly dendritic; 9-II-VI small, lateral, dorsolateral, or ventrolateral in po-

sition, removed cephalad a considerable distance from caudolateral angle; 9-

VII moderately to strongly developed, dorsal in position, removed cephalad a

considerable distance from caudolateral angle of tergite; 9-VIII strongly devel-

oped, ventral in position, removed cephalad a considerable distance from cau-

dolateral angle of sternite. Terminal Segments: hairs 1-IX, X absent; median

caudal lobe about half as long as broad; cercal lobe of female projecting only

short distance caudad of genital lobe, rounded apically; male genital lobe pro-

jecting slightly to far beyond median caudal lobe. Paddle: variously developed,

usually longer than broad, sometimes somewhat angular, sometimes emargi-

nate, never very narrow; external buttress more or less distinct and with short,

minute nearly invisible marginal spicules or almost smooth; midrib moderate-

ly developed; outer part of paddle larger than inner part; outer margin distad

of external buttress and inner margin usually smooth, without spicules; hair |

1-P present, single or branched, usually removed cephalad from margin of

paddle, often poorly developed and difficult to see; hair 2-P absent.

LARVAE. Head: varying from distinctly broader than long (phyllozoa) to

as long as broad; labrum short to moderately long (phyllozoa) and poorly dif-

ferentiated dorsally; mouthbrushes very numerous and filamentous; collar well

developed and narrow; ventral part of head capsule long; posterior tentorial

pit a considerable distance from caudal border; maxillary suture often not

complete anteriorly, straight or slightly concave mesally or laterally, and not

extending caudolaterad of tentorial pit; cephalic border of labial plate truncate

or concave anteriorly; aulaeum with long filiform spicules; mental plate well

developed and usually with 7-13 teeth on each side of large median tooth; chae-

totaxy quite uniform, complete except that hair 2-C is usually absent and hairs

16,17-C are always absent, hairs 4-7-C ordinarily multiple, 4 shortest, 5 usu-

ally longest and arising most caudad, and 7 with greatest number of branches

or hairs 4-7-C all strongly developed and subequal (flavithorax). Antenna: usu-

ally short to moderate in length; bare or with minute spicules (phyllozoa, fas-

cipes); chaetotaxy uniform, hair 1-A located in basal 0. 3-0.45, moderately de-

veloped, and usually 4-9b and hairs 2-6-A all terminal. Thorax: epidermal

pigment varying from absent to deep red or purple; integument usually not con-

Spicuously spiculose; spiracular sensilla distinct; notched organ not developed;

1 pair of conspicuous tracheal dilations present; all hairs except 13-P present;

hairs 9-12-P, M, T on common tubercles; 8-M, 7-T and sometimes 7-P with

basal tubercle; 1-3-P arising together, not close to midline; 4-7-P strongly

developed, 4 and 7 multiple, 5 and 6 usually single; 9-P multiple, 10,12-P sin-

gle, 12 usually somewhat longer; 14-P weakly developed; 1, 3-M moderately to

strongly developed; 2-M weakly developed; 4-M usually moderately developed;

5-7-M strongly developed, usually single; 8,9-M strongly developed, multiple;

10, 12-M strongly developed, single; 13, 14-M usually short and dendritic; 1-T

weakly to strongly developed; 2, 3-T usually moderately developed; 4-T multi-

ple, usually weakly to moderately developed; 5-T weakly developed; 6-T strong-

ly developed, single; 7,9-T strongly developed, multiple; 8-T usually dendri-

tic; 10,12-T single; 13-T multiple, moderately to strongly developed. Abdo-

men: pigmentation and spiculation as for thorax; spiracular sensilla distinct;

18 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

dorsal or more extensive sclerotized plate usually present on segment VII and

a smaller plate sometimes present on segment VI; hair 1 never palmate; 6-I, Il

strongly developed and multiple, 6-III, V, VI long and single, and 6-IV shorter,

single or double or only 6-I multiple and 6-II-VI double (arboricollis); 7-I mod-

erately to strongly developed, long, and single or multiple; 13-II, VI dendritic

or not (flavithorax); hairs 0, 8, 14-I and 14-II absent, otherwise all hairs pres-

ent. Segment VIII: mature larva with or without (alba) dorsal or more exten-

Sive sclerotized plate; comb scales usually in 2 rows, rarely in 1 row (albicos-

ta) or in 3 rows (anomalies); comb scales in posterior row usually larger than

those in anterior row; no comb scales ever attached to sclerotized plate. Si-

phon: moderately long to long; base of siphon often very irregularly sclerotized;

pecten absent; hair 1-S near middle or near base (alba, arboricollis), no acces-

sory hairs la-S; 2-S small, simple, always on siphon, no accessory hairs 2a-

S; valves small; spiracular apodeme distinct; trachea well developed. Anal

Segment: saddle usually complete; acus absent but sclerotized lateral band or

incomplete ring usually present basad of saddle; saddle margin without spic-

ules; hair 1-X on or caudad of saddle; 2-X very strongly developed, multiple;

3-X long, single; ventral brush well developed, usually with 6 (alba, arboricol-

lis, milloti), 7 (most species) or 8 (fascipes) pairs of hairs; grid well devel-

oped; no accessory saddle hairs; gills varied, dorsal usually longer than ven-

tral.

EGGS. Known only in Signifera group.

KARYOTYPE. Known only in Signifera group.

BIONOMICS. The immature stages of Orthopodomyia are found in plant

container habitats. While the majority of species breed in treeholes, others

breed in bamboo stumps and internodes, bromeliad leaf axils, and heliconia

flower spathes. In addition to the natural breeding sites, most species are oc-

casionally found in artificial containers. The larvae, which may live several

months, are filter feeders and apparently are not cannibalistic even when large

numbers occur in a small breeding site. |

The adults of all species are strictly sylvan and are rarely collected. Two

Species, albipes and andamanensis, are known to bite man and 3 species, alba,

kummi and signifera, are known to take avian blood. The feeding habits of the

remaining Species are unknown and the other activities of all species are un-

known.

SYSTEMATICS. Orthopodomyia is a small genus which is quite distinct

from all other mosquito genera in the adult and larval stages. Belkin (1962:

117) considered the genus to be distinctive enough to be worthy of recognition

as a separate tribe. The affinities of the genus are not evident, but similari-

ties with the genera Mansonia, Ficalbia and Culiseta have been noted. Edwards

(1932:106) felt that the genus was most closely related to Culiseta (as Theo-

baldia) and that the African treehole-breeding Culiseta (Theomyia) fraseri was

intermediate between the 2 genera.

The place of origin of the genus is difficult to determine. The Southeast

Asian group of species is the largest, most diverse, and apparently one of the

oldest groups, but the Middle American area contains 4 distinct, although

smaller and more homogeneous, groups of species.

Within the genus 8 distinct groups occur; these groups, with the number of

included species, are: Albipes, 9; Signifera, 6; Nkolbissonensis, 2; Thomasina,

2; Vernoni, 2; Albicosta, 1; Arboricollis, Ls and Folicolae, 1. Fach group is

largely or entirely restricted to one zoogeographic region. For the present I

am arranging the groups into 4 sections; when the immature stages of some of

Zavortink: Genus Orthopodomyia 19

the groups become better known it may be necessary to recognize additional

sections and it may be desirable to give the sections subgeneric status.

DISTRIBUTION. Orthopodomyia is largely confined to tropical and warm

temperate regions. The majority of species occur in Middle America and

Southeast Asia.

2(1).

3(2).

4(2).

5 (4).

6(5).

= 7(6).

KEYS TO GROUPS

Adults

(species 14 not included)

Tarsal segments 1 with yellow scales in distinct broad basal, central

and apical rings ..... . .ORTHOPODOMYIA SECTION, in part

wel eee NKOLBISSONENSIS GROUP

Tarsal deomenbe 1 without ae scales in distinct broad basal, central

dadiapicel Pings eee a ee a a ee oe ee

Mesonotum ornamented with 2-4 pairs of very narrow longitudinal lines

of pure white scales and broader bands of brown scales; wing without

light costal patches or broad sb hing He scales...

. BANCROFTIA SECTION 3

| Mesordhim mottled nah broad areas of light and dark scales or orna-

mented with 2 pairs of broad lines of silvery-white scales and bands

of brown scales; wing with conspicuous fea costal patches or broad

asymmetrical scales PEO ig he 8 a ers “ea LG ee eS

Mesonotum with 3 or 4 pairs of very narrow longitudinal lines of white

scales; hindtarsal tere 1 shorter than hindtibia; pra bristles 2-

ie ova. Se _ SIGNIFERA GROUP

Mesonotum 268 2 ee ai ie ae narrow | loneitudiaal dines of white scales;

hindtarsal sd as 1 OnEE? than hindtibia; pra bristles absent . .

Pee et es ae ; .ALBICOSTA GROUP ,

Parascutellar and pcx scale patches present; costal vein never with

more than inconspicuous basal, humeral and subcostal light patches .

: .THOMASINA SECTION

Parascutellar and pex scales normally absent: costal vein with more nu-

merous and more conspicuous light patches nit Bie aaa dead ate ar

Acrostichal scales almost entirely brown; pst with only a few scales

near ppl; ssp scales absent... ioe 66 -POLICOLAE, SECTION

Acrostichal scales largely light- colored: pst entirely covered with

scales; ssp scales present. . ORTHOPODOMYIA SECTION, in part 6

Costal vein lacking light patch opposite base of vein Rs (accessory sec-

toral patch); laterotergite with patch of light scales; acrostichal and

dorsocentral scales separated by longitudinal bare area. . .

, . ALBIPES GROUP

Costal v vein wath aceéssory sectoral light ‘patoh: laterotergite bare;

acrostichal and dorsocentral scales not separated by bare area .. 7

Lowermost mep scale patch small, rectangular and located close to

ail).

3(2).

4(3).

5 (4).

6(5).

7(6).

Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

upper patch; wing without broad asymmetrical scales; portion of cos-

tal vein along posterior edge of wing entirely dark- scaled. “eee

a . VERNONI GROUP

Lowermost m mep scale patch large, triangular and located in lower por-

tion of sclerite; wing with some broad asymmetrical scales; portion

of costal vein along posterior edge of wing with scattered light scales.

AO OE Bye . ARBORICOLLIS GROUP

Male Genitalia

(species 5 and 14 unknown)

Paraproct sclerotization with a single apical tooth; IX-T setae strong;

aedeagus acorn-shaped . . he NKOLBISSONENSIS GROUP

Paraproct sclerotization normally with at least 2 apical teeth; IX-T se-

tae fine or absent; aedeagus not acorn-shaped. ......... 2

Basal mesal lobe with 2 or 3 short, blunt and flattened stout bristles at

apex; VIII-T without a lobe on posterior Waren... «

‘THOMASINA SECTION

Basal mesal lobe with 3- 8 long, ‘pointed and terete stout bristles at apex;

VIII-T with a distinct lobe on posterior margin. .........3

Aedeagus more or less pyriform and spiniform broadened, flattened,

partially rolled and divided into numerous finger- -like lobes ieee

. SIGNIFERA GROUP

Without ‘both these. characteristics: aedeagus not pyriform or, if so, then

spiniform simple; spiniform not lobed or, if so, then lobes are few

and irregular and aedeagus is not pyriform ae ee Seago gtk,

Aedeagus with at least 15 small processes or teeth in a group or multi-

ple rows on each side of apex and sternal surface of aedeagus with 1

pair of strong processes or spines; sous with 2-4 irregular

lobes ... . . . ». VERNONI GROUP

Aedeagus without processes or r teeth at apex or, if so, then at most 9 in

1 row and sternal surface of aedeagus without strong processes or

Bpinee. Beri mie i on 8

Apex of aedeagus broadly truncate and with 5-8 crenate teeth on each

side and mesal surface of sidepiece with strong specialized bristles;

clasper with 2 spiniforms. . . . . . . .ALBICOSTA GROUP

Apex of aedeagus not broadly epunbate anid without teeth or, if narrowly

truncate and/or with pointed teeth, then mesal surface of Sidepiece

without specialized bristles; clasper normally with 1 spiniform . . 6

Apex of aedeagus with 2-7 large pointed teeth on each side; aedeagus

without a pair of strong teeth or spines on sternal wortade. “

‘ .FOLICOLAE SECTION

Apex of aedeagus without teeth: aedeagus often with a pair of strong

teeth or spines on sternal surface PT a ornare Sa ae NUR na eee ep

Aedeagus broadest in central portion and tapering towards both ends and

with a pair of sternal teeth near apex. . . . ARBORICOLLIS GROUP

2(1).

3(2).

4(1).

3(2).

4(3).

Zavortink: Genus Orthopodomyia 21

Aedeagus more or less pyriform or nearly as broad near apex as in cen-

tral portion or, if broadest in central portion, then sternal teeth not

NORT ANEK! FNS TEI ees eo eS COU

Pupae

(9. sampaioi, 11. vernoni, 12. milloti, 13. nkolbissonensis,

species 5 and species 14 unknown; 15. arboricollis not included)

Hairs 1-7-C all shorter than or subequal in length to trumpet and 6-C

cephalad of 7-C; 1-VI usually mesad of 2-VI. ... . 2

At least hairs 5-C and/or 7-C not less than 1.5 times length ot trumpet ;

or 6-C caudad of 7-C; .1-VI laterad of 2-VI° .°. 202. . 4

Hair 3-C subequal in length and number of branches to 1-C; surface of

paddle smooth or with minute ia phe se in posterior portion of in-

ner part near inner margin . . . . . . SIGNIFERA GROUP

Hair 3-C much shorter and usually with t more numerous branches than

1-C; entire surface of paddle with minute spicules. ....... 3

Hair 1-II 3-5b; external buttress of paddle extending only slightly distad

of level of apex of median caudal lobe. . . . . .ALBICOSTA GROUP

Hair 1-II usually 10-13b (6-16); external buttress of paddle extending far

distad of level of apex of median caudal lobe. .THOMASINA SECTION

Hair 3-C weakly developed, not as strong as 2-C; 5-C most strongly de-

veloped hair on cephalothorax; 9-VII moderately developed, usually

3-6b and not more than 0. 33 length of 9- VIII. .FOLICOLAE SECTION

Hair 3-C weakly to strongly developed, always at least as strong as 2-C;

7-C usually most strongly developed hair on cephalothorax; 9-VII

ne Nie: 8-12b iis ae and about 0.5-0.75 length of 9-VIII

: ; . ALBIPES GROUP

Larvae

(13. nkolbissonensis and species 14 unknown;

9. sampaioi and 11. vernoni not included)

Comb scales normally inl row... .%. .... ...ALBICOSTA GROUP

Compiseales normaly IE Wow ere ee es Vee ee

. Hair 6-II-VI similarly developed, double; 2-III-VII stellate; basal por- —

tion of siphon and anal saddle with spinules . ARBORICOLLIS GROUP

Hair 6-II developed like 6-I and usually more than double, 6-II-VI nor-

mally single or only 6-IV double; 2-III-VII usually small and sein

never stellate; siphon and anal saddle without spinules . j

Hairs 4-10-C all more or less equally developed, long, strong and mul-

tiple; 0,1, 3-7-P all more or less equally developed, long, strong and

multiple fee) a a pe ate eit ¢ Adee Se 1 Dart

ghd gt cele Gea tate) i teh flavithorax

Hairs 4-10-C not all equally developed, in n particular @ 5, Ue. oe 1-3b;

0,1, 3-7-P not all equally developed. ‘ es : 4

Hairs 12,13-I and 1, 5-VIII stellate; ventral brush with 6 pairs of hairs .

22 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

. . .VERNONI GROUP

Hairs 12, 13- I and 1, 5- Vl not stellate: ventral ‘brush wit with 7 or 8

pairs of WAS Ue ae a Tas Nt ee ee)

5(4). Hair 4-M double or multiple; 2-VI, VII mesad of hair 1 of corresponding |

segment... 6

Hair 4-M single; 2- VI, VI laterad of hair t of corresponding segment. it

6(5). Comb scales in posterior row usually 2-4 (1-6); hair 13-I more strongly

developed than 10-I; 2-I, II mesad of hair 1 of corresponding segment

; .FOLICOLAE SECTION

Comb scales in n posterior row ‘usually 7- 12 (5- 14); hair 13-I less strong-

ly developed than 10-I; 2-I, II laterad of hair 1 of corresponding seg-

Mel Keith Se oe eo . . . . . ALBIPES GROUP, in part

7(5). Ventral brush with 8 pairs of hairs, the strongest hairs with about 16-23

branches; hair 5-IV-VI usually longer than hair 3 of corresponding

segment; head capsule and antenna with minute spicules .. .

oe -THOMASINA SECTION

Ventral. brush usually eit 6 or 7 pains: of hairs, the strongest hairs with

about 4-12 branches; hair 5-IV-VI usually shorter than hair 3 of cor-

responding segment; head capsule and antenna without spicules . .

Rite hs ees . SIGNIFERA GROUP

BANCROFTIA SECTION

FEMALES. Head: eyes partially or completely separated above antennae,

or entirely contiguous (albicosta); frontal and orbital bristles dark; ocular bor-

der present, white, consisting of narrow curved decumbent scales passing cau-

dad of lateral orbital bristles and cephalad of mesal ones; vertex and occiput

with few to numerous decumbent scales, all narrow and curved or with some

broad and flat (albicosta), and numerous erect scales, cephalad ones longer and

with fewer forks; clypeus an elongate angular lobe or triangular (albicosta,

Kummi subgroup), usually bare; labium considerably swollen apically, weakly

developed shaft bristles present near apex; palpus straight, 0.33 to 0. 48 of

proboscis, 4- or 5- (pulchripalpis) segmented; torus with scales; first flagel-

lar segment not or slightly (albicosta) swollen; no (albicosta), 1 (pulchripalpis)

or several proximal flagellar segments with scales. Thorax: paratergite nar-

row to moderately broad, bare; posterior dorsocentral bristles sometimes in

a double row; posterior fossal bristles rarely absent (albicosta); fossal bris-

tles present or absent; all meSonotal and scutellar bristles dark; mesonotal

ornamentation consisting of 4-6 pairs of lines of small white scales arranged

into pattern of 2-4 longitudinal lines and broader areas or lines with brown

scales; parascutellum normally bare; lateral lobes of scutellum with or without

scales; pleural bristles, except as noted below, dark; upper anterior apn bris-

tles 1-8 strongly developed, 0-4 weakly to moderately developed, lower apn

bristles 0-2 strongly developed, 0-4 weakly to moderately developed, at least

1 bristle always present, sometimes light in color; ppl bristles 4-8 strongly

developed, 0-9 weakly to moderately developed; pra bristles absent (albicosta)

or 2-12; upper stp bristles 1 or 2, moderately to strongly developed, posterior

stp bristles 2-10, weakly to strongly developed, lower stp bristles 0-10, weakly

Zavortink: Genus Orthopodomyia 23

to moderately developed; upper posterior mep bristles 4-15, sometimes light

in color; anterior mep bristles rarely present (pulchripalpis); pleural scaling

consisting of 3 more or less distinct horizontal lines and 1 nearly vertical line

of white scales. Legs: coxal bristles light or dark; hindtarsal segment 1

shorter or longer (albicosta) than hindtibia; hindclaws smaller than fore- and

midclaws. Wing: remigial bristles present, weakly developed; cell R9 varied

in length, vein R2 1.75-3.50 of vein R943; cell M9 varied in length, vein

M1+2 1.20-2.50 of vein M distad of crossvein m-cu; plical area with a few

light scales at base ventrally; alula scales narrow; upper calypter with bris-

tles; scales symmetrical, or nearly so, varying in shape and degree of spread-

ing; ornamentation varied, but without costal spots or yellow or golden scales.

Abdomen: tergite I with middorsal fan of scales and sometimes with scattered

white scales, laterotergite bare.

FEMALE GENITALIA. Tergite IX with or without dorsolateral setae.

MALES. Essentially as infemales. Labium: not more strongly swollen

than in females. Palpus: 0.77-1.02 of proboscis, segment 3 with or without

well developed apical bristles. Antenna: torus greatly enlarged; flagellum

strongly plumose; segments 12 and 13 subequal in length; segment 1 with or

without (albicosta) scales. Legs: anterior fore- and midclaws with basal tooth

too small to be seen on pinned specimens, posterior fore- and midclaws uni-

serrate with tooth too small to be seen on pinned specimens. Abdomen: side-

piece with light and dark scales.

MALE GENITALIA, PUPAE, LARVAE. Varied. See descriptions for Sig-

nifera group and albicosta.

BIONOMICS. The immature stages are found in bamboo internodes, tree-

holes and artificial containers. Females are not known to bite man.

SYSTEMATICS. The Bancroftia section is made up of the Albicosta and

Signifera groups. The former group contains only the South American albicos-

ta while the latter, centered in North America, contains 6 species. I am com-

bining these groups because the adults of both share a highly derived ornamen-

tation. However, the male genitalia, pupa and larva of albicosta are no more

Similar to the Signifera group than to any other group.

This section appears to have arisen in the American tropics and to have

split into the South American albicosta and the North American centered Signi-

fera group early in its history. The southern-most species in the Signifera

group, kummi, is somewhat intermediate in adult ornamentation between albi-

costa and the other members of the Signifera group.

The affinities of this section are not obvious. The male genitalia of albi-

costa, particularly in the shape and dentition of the aedeagus, are reminiscent

of the Madagascan Vernoni group. The male genitalia of the Signifera group

resemble the Vernoni group and the Thomasina section in the shape of the spin-

iform and are like some species in the Oriental Albipes group in gross aedea-

gus Shape. Among the groups for which the pupa is known, the pupa of both the

Albicosta and Signifera groups is most similar to that of the Thomasina section.

In fact, the pupa of albicosta is somewhat intermediate between that of the Sig-

nifera group and the Thomasina section. The larva of albicosta shares charac-

teristics with the middle American phyllozoa and with the Signifera group. The

larvae in the Signifera group bear a close resemblance to some species in the

Albipes group (especially the Anopheloides subgroup) and the Thomasina sec-

tion.

DISTRIBUTION. North America to southern Europe, south to South Amer-

ica and North Africa.

24 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

KEYS TO GROUPS

See keys on p. 19-22.

SIGNIFERA GROUP

FEMALES. Head: eyes partially to completely separated above antennae

by white decumbent scales, which may be accompanied by weakly developed

dark bristles; orbital bristles usually 3, 4 lateral pairs and 4, 5 mesal pairs;

vertex and occiput with few to numerous white curved decumbent scales ar-

ranged into a more or less anteriorly directed "'V''-shaped pattern, and brown

or black erect scales; dorsolateral broad scales all white (Signifera subgroup)

or white and dark (pulchripalpis and Kummi subgroup); lateral and ventral

scales white; suborbital bristles dark; clypeus an elongate angular lobe or

triangular (Kummi subgroup), usually bare; palpus 0. 33 to 0. 48 of probiscis,

4- or 5- (pulchripalpis) segmented, segments 4 and 5 entirely white-scaled

dorsally; torus with dorsomesal small white flat scales; flagellum with seg-

ment 1 not swollen, segment 1 with brown scales dorsomesally (pulchripalpis)

or segments 1-3, 4,5,6, or 7 with dorsomesal line of white scales; dorsal fla-

gellar bristles 2.0-2.5 times length of their segment on proximal segments,

ventral bristles usually much reduced or absent on segment 1, usually some-

what reduced on 1 to several succeeding segments. Thorax: paratergite nar-

row to moderately broad; humeral, lateral prescutal and posterior fossal bris-

tles usually numerous enough to completely enclose fossa; mesonotal ornamen-

tation consisting of 4-6 pairs of narrow lines of small white scales, arranged

into a pattern of 3 or 4 pairs of longitudinal lines, and broader areas or bands

with very small brown scales; white scales arranged as follows: (1) lateral

prescutal scales always present, in line from anterior promontory to scutal

suture laterad of humeral and lateral prescutal bristles, (2) supraalar scales

present in line from scutal suture to parascutellum mesad of supraalar bris-

tles or absent (signifera, in part), (3) inner dorsocentral scales always pres-

ent, in line from anterior promontory to prescutellar space laterad of acros-

tichal bristles, (4) lateral prescutellar scales always present, larger than other

mesonotal scales, in line from anterior edge of prescutellar space to scutellum

mesad of lateral prescutellar bristles, rarely connected anteriorly to inner

dorsocentral scale line (waverleyi), (5) posterior fossal scales rarely present

(waverleyi), scattered in posterior fossal area and connected to outer dorso-

central scale line, (6) outer dorsocentral scales always present, in line from

scutal suture to about level of cephalic end of lateral prescutellar scale line or

extending to scutellum (pulchripalpis), laterad of posterior dorsocentral bris-

tles; brown scales arranged as follows: (1) in band accompanying but extending

further caudad than acrostichal bristles, (2) in band accompanying dorsocentral

bristles, (3) as sparse covering in all or part of fossal area; lateral lobes of

scutellum with or without scales, midlobe with 1 pair of narrow white scale

lines which extend 1 to 2 times length of scutellum from its caudal margin; up-

per anterior apn bristles 2-8 strongly developed, 0-4 weakly to moderately de-

veloped, lower apn bristles 0-2 strongly developed, 0-4 weakly to moderately

developed; ppl bristles 3-8 strongly developed, 0-9 weakly to moderately de-

veloped; pra bristles 2-12; upper stp bristles 1 or 2, moderately to strongly

developed, posterior stp bristles 3-10, weakly to strongly developed, lower stp

bristles 0-10, weakly to moderately developed; upper posterior mep bristles

4-15, not always dark in color, anterior mep bristles rarely present (pulchri-

Zavortink: Genus Orthopodomyia 29

palpis); pleural scaling with individual lines narrow (Kummi subgroup) or broad-

er; apn with diagonal row of narrow curved decumbent scales between two

groups of bristles and occasionally with 1, 2 similar white scales above upper

anterior bristles; ppn with narrow horizontal row of flat scales extending from

line on apn to its caudal border ventrad of posterior ppn bristles or with scales

reduced or absent (signifera, in part); psp bare or with scales in dorsal poste-

rior patch; pra usually with few to many curved decumbent scales; ppl with

diagonal row of broad flat scales below ppl bristles, sometimes with a few sim-

ilar scales above bristles; pst bare, with broad flat scales near ppl (pulchri-

palpis), or in horizontal line across entire prosternum (Kummi subgroup); pcx,

ssp and hypostigial scales absent; stp scaling variable, with 3 more or less

distinct lines of scales, as follows, broad flat scales in 2 horizontal lines, 1

above and 1 below upper stp bristle, and broad flat and sometimes semierect

scales in nearly vertical line extending ventrad from level of lower margin of

mep along posterior stp margin, both horizontal lines sometimes joined before

or behind upper stp bristle or vertical line broad and extending dorsad over

both horizontal lines; mep with upper diagonal patch or line of broad semierect

scales, some elongated, and lower anterior patch or line of broad flat scales,

lower patch sometimes joining upper or rarely absent or much reduced (signi-

fera, in part); a few additional large brownish or transparent scales may be

scattered over pleuron, especially meron and stp, of freshly eclosed, robust

adults. Legs: coxal bristles usually very dark; broad flat scales present along

outer edge of anterior surface on all coxae and above and behind uppermost bris-

tles on forecoxa; trochanters with flat scales on anterior, inner and posterior

surfaces; femora usually with small white knee spots; tibiae with apical white

patch if tarsal segment 1 is white at base; hindtarsal segment 1 shorter than

hindtibia; hindtarsal segment 1 with medium-sized posteriorly incomplete white

ring basally and complete or ventrally incomplete white ring apically, segments

2-4 usually with conspicuous dorsal white patches at both ends, those on proxi-

mal segments sometimes forming complete rings, segment 5 entirely white-

scaled dorsally or all or partially dark (waverleyi and part of signifera); patch

or ring at joint between segments 1-2 largest, that over each succeeding joint

smaller. Haltere: scales whitish. Abdomen: tergite I fan white-scaled; ster-

nite VIII largely bare, a few scales usually present laterally.

FEMALE GENITALIA. Tergite IX with dorsolateral setae.

MALES. Essentially as infemales. Palpus: 0.93-1.02 of proboscis, seg-

ment 5 small, segment 3 with or without strong apical bristles; white scales in

patch at base of segment 4 and covering all of segment 5. Antenna: integument

of flagellum dark except for lighter bands basad of whorls on shorter segments;

segment 1 with dark or dark and white scales dorsomesally; bristles of flagel-

lar whorls about 0.4-0.5 length of entire flagellum on proximal segments, about

0.5-0.8 length of segment 13 on that segment. Wing: plical scales often absent.

MALE GENITALIA. Segment VIII: tergite with distinct lobe on posterior

margin. Segment IX: tergite with 0-6 fine bristles on each side. Sidepiece:

medium to long conical; basal mesal lobe with 3-7 terete apical bristles, tergal

2-6 quite stout, sternal 1-3 finer; mesal surface membranous for short dis-

tance distad of basal mesal lobe; mesal surface without specialized setae dis- —

tad of basal mesal lobe; tergomesal surface with 1 or 2 very large, and usually

1 or 2 smaller, curved, specialized bristles distad of or near basal mesal lobe.

Spiniform: 1, apex flattened, broadened, partially rolled and divided into nu-

merous more or less regular finger-like lobes. Aedeagus: strongly sclerotized,

more or less pyriform, with 1 pair of triangular spines sternally and 1-7 small

irregular processes on an apical ridge tergally. Proctiger: paraproct sclero-

tization with 2-5 apical teeth; cercal setae 2-6.

26 : Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

PUPAE. Cephalothorax: lightly to moderately pigmented, with upper and

posterior portions of wing case and portion of mesonotum caudad of trumpet

darker; integumentary sculpturing absent to very weak, reticulate; smaller

hairs concolorous with integument, larger hairs darker; hairs 1-3-C all more

or less equally developed, weak to moderate; 4,5, 7-C moderately and more or

less equally developed; 6-C cephalad of 7; 8,9-C sometimes strongly devel-

oped. Trumpet: deeply pigmented except at extreme base and apex and much

darker than darkest part of cephalothorax or more or less uniformly moderate-

ly pigmented and not darker than darkest part of cephalothorax (pulchripalpis);

distal portion gradually widened from base; pinna medium-sized. Metanotum:

lightly to moderately pigmented, with haltere cases lighter; integumentary

seulpturing absent or very weak, reticulate; never more than 3 pairs of hairs

developed. Abdomen: lightly to moderately pigmented; when moderately pig-

mented posterior portion of anterior segments and posterior segments in gen-

eral are lighter; segments II-VHI usually with weakly to moderately developed

reticulate to imbricate or somewhat spiculate integumentary sculpturing; small-

er hairs and branches concolorous with integument, larger hairs darker; hair

1-II not dendritic and usually 2, 3b (1-5) or dendritic and usually 4-6b (4-16)

(signifera, in part); 2-II at level of or caudad of 3-II; 1-VI, VH mesad of hair

2 of corresponding segment; 1-VI usually moderately long and 2, 3b; 9-VII

moderately developed, usually 3-7b, longest branch shorter than or only slight-

ly longer than 0.5 distance between alveoli of hairs 9-VII and 9-VIH; 9-VIII

strongly developed, usually 7-10b (6-11), longest branch seldom extending

- much beyond end of external buttress of paddle. Terminal Segments: male gen-

ital lobe projecting considerably beyond median caudal lobe. Paddle: lightly to

moderately pigmented, midrib darker; longer than broad, shape varied; ex-

ternal buttress long, more or less straight in middle portion, nearly smooth;

midrib usually convex mesally; surface of paddle smooth or with a band of

spicules near inner margin (kummi).

LARVAE. Head: slightly broader than long to as long as broad; integu-

ment without ornamentation; labrum short; hair 2-C sometimes present. An-

tenna: short; integument smooth. Thorax: hair 4-M always single. Abdomen:

segment VI often with a sclerotized dorsal plate and segment VH usually with a

very large sclerotized plate (both plates always absent in alba); hairs 2-I, Il

laterad of hair 1 of corresponding segment; 6-I, II usually multiple (sometimes

only double in alba), 6-III-VII single; 13-I shorter than 10-I; 5-IV-VI shorter

than hair 3 of corresponding segment; 2-VI, VII laterad of hair 1 of correspond-

ing segment. Segment VIII: large sclerotized plate present or absent (alba);

comb scales in 2 rows; posterior row usually with 5-8 (5-12) comb scales;

posterior comb scales with 1 long apical spine and basal fringe. Siphon: integ-

umentary sculpturing absent or weakly developed, imbricate. Anal Segment:

saddle usually complete, Sometimes ventrally incomplete in alba; sclerotized

band or incomplete ring present basad of saddle; integumentary sculpturing as

for siphon; ventral brush usually with 6 pairs of hairs in alba, usually with 7

pairs of hairs in other species, the more caudal hairs usually with 4-8 branches

in alba and 8-12 branches in other species.

EGGS. Length: about 0.61 mm. Maximum Width: about 0.14 mm. Laid

Singly or in small groups above the water meniscus on sides of breeding site;

not resistarit; brown to black in color; cephalic end larger and rounded, caudal

end narrower and tapering; morphologically ventral surface uppermost in laid

egg and with a veined gelatinous veil; hatching suture oblique, inclined about

- 45° from longitudinal axis of egg.

KARYOTYPE. Breland (1961:367-368) reported that a pair of satellites

was associated with I of the larger pairs of chromosomes during prophase in

Zavortink: Genus Orthopodomyia 27

the prepupal brain of both alba and signifera. The members of the pair of sat-

ellites were sometimes unequal in size and it was postulated that they may rep-

resent sex chromosomes.

BIONOMICS. With the exception of kummi, the species in this group are

restricted to breeding in treeholes and artificial containers. Kummi not only

occupies these habitats, but also is found in bamboo stumps. Females of some

Species are known to take avian blood; no species is known to bite man.

SYSTEMATICS. This is the most recent and highly derived group in the

genus Orthopodomyia. It contains 6 very closely related species: alba, kummi,

pulchripalpis, signifera, waverleyi, and species 5. With the exception of alba

and signifera, which are sympatric over much of the eastern United States, the

species are allopatric and form a superspecies. The species in this group are

the most poorly marked of any in the genus. The male genitalia are almost

identical in all species; barring alba and possibly species 5, the larvae cannot

be adequately separated (see below) and separation of the pupae, while some-

what better, is still rather poor. The adults of all species except alba and sig-

nifera are easily separated on the basis of ornamentation. I am considering the

allopatric forms in this group to be species rather than subspecies because 2

of them, signifera and kummi, meet in southeastern Arizona without extensive

hybridization and the differences between most of the other forms are of the

Same magnitude as those between kummi and Signifera.

Larval features have historically been used for the recognition and separa-

tion of the forms (especially alba, californica and signifera) of this group which

occur in the United States (see Bohart, 1950:399); Carpenter and LaCasse,

1955:98; Chapman, 1965:438). However, I have largely rejected the hypothesis

that larval characters are important for separating species in this group. This

decision is based on considerable study of the adults and the complete larval

and pupal chaetotaxy of all the forms inthe Signifera group. Indeed, if one

were to base North American species on larval characters alone, several addi-

tional species would have to be recognized. More specifically, at least 4 more

larval species with signifera-like adults and 1 with a kummi-like adult would

be described. Moreover, the 2 larval species with kummi-like adults would

scarcely be separable from 2 of the larval species with signifera-like adults.

In addition, if one considers pupal characters to be as important as larval, then

3 additional species, 2 with signifera-like adults and 1 with a kummi-like adult,

would be recognized. Many of the resulting 12 ''species" with either signifera-

like or kummi-like adults would have disjunct distributions and some areas

would have several species occurring sympatrically. One would repeatedly

need to postulate hybridization as an explanation for the mixture of imma-

ture and adult characters in the different larval or adult lines. In short, if

Species are based on larval and pupal chaetotaxy, then species are obtained

which have biologically unsound distributions and any attempt to explain the

evolution of the group must be highly speculative.

On the other hand, in a classification based largely on adult ornamentation,

only 6 species are recognizable in the entire Signifera group. Such an approach

is suggested when one considers that the three forms in the Signifera group

which differ most in adult ornamentation, and which are most likely to be spe-

cifically distinct on the basis of geography (the Central American kummi, the

North American signifera and the European pulchripalpis), have nearly identi-

cal larvae. The 6 species delimited under such a system have continuous dis-

tributions and all of the species except alba and signifera are allopatric. Ex-

planation of the evolution of the group need not be highly speculative, since all

Species could have arisen in geographical isolation. In summary, if species

28 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

are based largely on adult ornamentation, then the resulting species have bio-

logically sound distributions and the explanation of the evolution of the group is

simple. This approach seems more reasonable than the one previously outlined

and it is the one followed here. Under this system 2 of the 6 species, signifera

and kummi, have various non-hairy, semi-hairy and hairy larval and pupal

forms and only 1 of the 6 species, alba, is Sey known to have a diagnostic

larva.

The Signifera group can be divided into 3 subgroups on the basis of adult

ornamentation; the Kummi subgroup contains kummi and species 9, pulchripal-

pis forms a monotypic subgroup, and alba, signifera and waverleyi form the

Signifera subgroup.

I believe the ancestral species in this group arose in the Middle American

area and was derived from the same stock as albicosta. Since the adult wing

and pleural ornamentation of kummi are similar to albicosta and kummi is the

most southern species in the group, I am considering kummi to be the most |

primitive species.

Although I am including keys to the male genitalia, pupae and larvae of the

species in this group, as indicated above, positive identification of most species

can be made only on the basis of adult ornamentation.

DISTRIBUTION. United States to southern Europe, south to Panama, the

Greater Antilles, and North Africa.

KEYS TO SPECIES

Adults

i. Outer dorsocentral scale line extending to scutellum; scales on.vein 1A

very narrow and lying parallel with vein; line of white scales at base

of vein R about twice length of remigium . .

as (PULCHRIPALPIS SUBGROUP) 4. pulchripalpis

Giter dorsocentral scale line not extending to scutellum; scales on vein

1A narrow to broad, Spreading; line of white scales at base of vein R

restricted to remigium or extending to Separation of veinRg ... 2

2(1). Pst with a complete horizontal line of white scales; scale lines of pleu-

ron very narrow; base of vein 1A dark-scaled. . .

: (KUMMI SUBGROUP) 3

Pst Pinout scales: scale ‘dines 6 on pleuron t broader; base of vein 1A

Tete Bee ee er (SIGNIFERA SUBGROUP) 4

3(2). Line of white scales at base of vein R restricted to remigium .Species 5

Line of white scales at base of vein R extending to separation of vein Rg

6. kummi

-4(2). Lateral prescutellar scale line connected to inner dorsocentral scale

line; posterior fossal white scales present and connected to sparsely

scaled outer dorsocentral scale line ... . . 1. waverleyi

Lateral prescutellar scale line not connected fo" inner dorsocentral scale

line; posterior fossal white scales absent and outer dorsocentral

scale line densely scaled ee ee ee iy ate, ok a ee a

5(4). Lower stp bristles usually 4-10 in number and present throughout length

e(1).

3(2).

4(3).

Zavortink: Genus Orthopodomyia 29

of lower stp scales; base of vein R4+5 with patch of white scales, or

if not, then entire white fs over crossveins is much reduced.

yi rae signifera

Lower cab bristles ‘Aeually 0- 2 in 1 number and present only in uppermost

portion of lower stp scales; base of vein R4+5 usually dark-scaled

and white spot over crossveins VOTES eee Rc Chal! Bide LOR

Male Genitalia

(Species 5 unknown)

Clasper not constricted distad of insertion of apodeme, but tapering

smoothly from base to apex; strong setae of basal mesal lobe usually

ALT SEPAIERES is) dos M0 . « « «4. pulchripalpis

Clasper usually constricted ‘distad of iisertion of apodeme, so that mid-

dle portion is narrower than apical portion; strong setae of basal

mesal lobe usually curved mesally at gh

.1. waverleyi; 2. signifera: 3. alba: 6. “kummi

Pupae

(Species 5 unknown)

Paddle with few to many minute spicules in posterior portion of inner

part near inner margin; hair 5-IV, V almost always extending to cau-

dal margin of following segment; 6, 7-I strongly developed, long.

d 6. kummi

Paddle without mirube spicules: hair 5 IV, V usually not extending to

caudal margin of following segment; 6, 7-I weakly to strongly devel-

oped, short to long . hte Paes Fie geal Ge

Hair 2-II-VI 0.4-0.6 length of hair 1 of corresponding segment; 5-III

similar in development to and as, or nearly as, strongly developed

as 3-III and 5-IV-VI; 5-IV, V not extending beyond 0.6 of following

segment; 6, 7-I weakly developed, short. ... ia nee Wale

Hair 2-II-VI less than 0.3 length of hair 1 of corresponding segment;

o-III not as strongly developed as 3-III and 5-IV-VI; 5-IV, V variable

in length; 6, 7-I weakly to strongly developed, short to long hg a

Trumpet amber to tan colored and concolorous with or lighter than dark-

est portion of cephalothoracic integument; hair 5-IV, V not extending

beyond 0.8 of following segment; 6, 7-I weakly to moderately devel-

oped, short to moderate in length. .. . - a se @, DUICNTIpelpis

Trumpet deep tan to brown and darker than darkest portion of cephalo-

thoracic integument; hair 5-IV, V variable in ie lid fo | Puasa to

strongly developed, shortto long. ..... 4.

Hair 6-I, II not as strongly developed or as long as hair 3 of correspond-

ing segment and paddle index 1.45-1.65; 5-IV, V not extending beyond

0.6 of following segment. . ... . .1. waverleyi; 2. some signifera

Hair 6-I, II at least as strongly developed and as long as hair 3 of cor-

responding segment or, if not, then paddle index 1.15-1.45; 5-IV, V

variable in length, sometimes extending to caudal margin of following

ric ai nl ag near Ga MP EAN UR MeAEDe ym GL Ula a Mera rane rie. | hd «ied ya

30 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

Larvae

7. Hair 1-S weakly to moderately developed, usually 3, 4b (2-6), arising

near base of siphon, and with branches usually not extending much

beyond middle of siphon; 3-VIII weakly to moderately developed, usu-

ally 2, 3b (1-5), and usually shorter than the longest comb scales;

13-C strongly developed, usually 7,8b (5-12) ...... . .3. alba

Hair 1-S strongly developed, usually with 6 or more branches, arising

near middle of siphon, and with branches extending nearly to or be-

yond apex of siphon; 3-VIII strongly developed, usually with 5 or

more branches, and at least 1.5 times length of longest comb scales;

13-C moderately developed, usually 2, 3b (1-4). .........2

2(1). Hair 13-T very strongly developed, 6-11b, and as long as 6-II; 6-II 10-

14D 3 oo. . Species 5

Hair 13-T moderately is strongly dey eloped: usually 2- 4b (1- pe and

usually not nearly as long as 6-II; 6-II usually 5-7b (2-8) . eee

3(2). Hair 1-T longer than 2-T; 1-II often at least 1.5 times length of 3-II .

. 2. Ssemi-hairy andhairy forms of signifera; 6. hairy forms of kummi

Hair 1-T shorter than 2-T; 1-II usually subequal to or shorter than 3-II

4(3). Siphon index usually greater than 4.0; diameter of base of anal saddle

not or only slightly smaller than diameter at most distal point along

ventral surface, so that dorsal and ventral surfaces of saddle appear

parallel when viewed from the side; hair 13-VII usually 5, 6b (5-8).

ue 4, pulchripalpis

Sinhon anew eur less a 3 6: diameter of base of anal saddle usu-

ally smaller than diameter at most distal point along ventral surface,

so that dorsal and ventral surfaces diverge distally when viewed from

the side; hair 13-VIt often only 3,40 (226). ee ee 8

5(4). Branches of hair 1-S extending beyond apex of siphon and dorsal anal

gills 1.1-2.1 length of anal saddle . . 2. signifera, in part; 6. kummi

Branches of hair 1-S not extending beyond apex of siphon, or, if so, then

dorsal anal ati — 0.45-0. 80 length of anal saddle. ;

: ; ; Lae ss .l. waverleyi; 2. signifera, in 1 part

SIGNIFERA SUBGROUP

FEMALES. Head: vertex and occiput with white decumbent scales numer-

ous, evenly distributed; dorsolateral broad scales all white; clypeus an elon- ©

gate angular lobe, usually bare; labium dark-scaled with numerous white

scales in distinct line or line of speckles from base to or near apex on each

side of labial groove; palpus 0. 33 to 0.35 of proboscis, 4-segmented, segment

4 very short, segments 1, 2 and 3 dark-scaled with white scales in line or line

of speckles along dorsal surface; flagellar segments 1-3, 4, 5,6 or 7 with white

scales. Thorax: paratergite moderately broad; outer dorsocentral scale line

not extending to scutellum; lateral lobes of scutellum with or without scales;

anterior mep bristles absent; psp scales present or absent; pst without scales.

Zavortink: Genus Orthopodomyia 31

Legs: coxae with white scales or scine lower ones on forecoxa brown or black;

trochanters with white or white and dark scales; forefemur with anterior sur-

face usually streaked with white scales in basal 0.05-0.6 dorsally, remainder

dark-scaled with white speckles, ventral margin speckled or lined with white

scales from base to apex, posterior surface usually with narrow ventral dark

streak from base to apex, area dorsad of streak largely white-scaled basally

and white-scaled with dark speckles apically; midfemur with anterior surface

largely dark-scaled with white speckles, base varied, posterior surface pale-

or white-scaled in basal 0.05-0.5 and to or near apex in progressively narrow-

ing streak near ventral margin, remainder dark-scaled with white speckles;

hindfemur with anterior surface usually pale- or white-scaled in basal 0. 05-

0.4, and somewhat further distally along ventral margin, remainder dark- scaled

with white speckles, posterior surface like that of midfemur except that greater

proportion of base is usually light-scaled and light scales do not extend to apex;

tibiae largely dark-scaled; foretibial anterior surface with white scales in line

or line of speckles from base to or near apex, posterior surface usually with

similar but often shorter and/or less well defined line; midtibial anterior sur-

face with white scales usually in poorly defined line or line of speckles from

base to or near apex, posterior surface with well defined white line, ventral

surface with white speckles basally; hindtibia varied; tarsi largely dark-scaled,

varied. Wing: speckled and spotted with many white scales, vein R with remi-

gium white-scaled, base of vein 1A white-scaled to about level of base of vein

Rg, area of crossveins usually with more or less conspicuous white spot made

up of white patches on veins Rg, M, and Cu, base of veins Rg and M often white-

scaled, remainder of veins mentioned and all other veins usually dark-scaled

with numerous white speckles; veins behind vein Rj with broad scales; scales

on vein 1A wide-spreading. Haltere: scales broad.

MALES. Essentially as infemales. Palpus: 0.93 to 1.02 of proboscis,

segment 3 with or without well developed apical bristles; segments 1, 2 and 3

with white scales in line or line of speckles on dorsal surface. Antenna: flagel-

lar segment 1 with white and dark scales dorsomesally. Thorax: upper poste-

rior mep bristles reduced in number. Wing: white-scaling generally reduced;

seales at base of vein 1A not wide-spreading.

MALE GENITALIA. Segment IX: sternite without small projection in mid-

dle of cephalic edge. Clasper: usually strongly constricted distad of origin of

apodeme so that middle portion is narrower than distal portion. Aedeagus: bas-

al portion usually rounded, sometimes slightly angular.

SYSTEMATICS. This is the most highly derived subgroup in the Signifera

group. It contains 3 species, alba, signifera and waverleyi. Signifera, the

widespread, dominant species, is found throughout most of the United States,

in northern Mexico and on all the Greater Antilles except Jamaica; waverleyi

is restricted to Jamaica; alba is found throughout the eastern United States and

in northeastern Mexico. Alba is apparently the most primitive of the 3 species.

DISTRIBUTION. United States, south to northern Mexico and the Greater

Antilles.

1. Orthopodomyia waverleyi (Grabham)

Figs. 2, 3,4

1907. Mansonia waverleyi Grabham, 1907:25. *TYPE: Lectotype & with geni-

talia slide 650626-14, specimen bearing the handwritten label "'signifer,

32 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

Jamaica, Dr. Grabham, 1.9.06,'' Waverley Estate, Constant Spring,

[St. Andrew, Surrey], Jamaica, reared from a larva found in a mango

treehole, 1 Sept 1906, M. Grabham; PRESENT SELECTION [USNM].

Reduced to synonymy with signifera by Dyar (1928:397); resurrected

from synonymy with signifera by Edwards (1939:121, 123); reduced again

to synonymy with signifera by all subsequent workers: resurrected from

synonymy with signifera in present study.

Orthopodomyia waverleyi of Howard, Dyar and Knab (1917:891-893); Johnson

(1919:424); Bonne and Bonne-Wepster (1925:488); Gowdey (1926:74); Root

(1927:463, 465, questionable identification); Edwards (1939:121, 123).

Pneumaculex waverleyi of Theobald (1910b:469, 619-620).

Orthopodomyia signifera of Dyar (1928:397, in part): Edwards (1932:108, in

part); Lane (1939:97, in part); Hill and Hill (1945:3); Thompson (1947: 79);

Hill and Hill (1948: 51): Lane (1953:628-629, in part); Stone, Knight and

Starcke (1959:124, in part); Stone, Sabrosky, Wirth, Foote and Coulson

(1965:109, in part).

FEMALE (fig. 2). Wing: 3.78 mm. Proboscis: 2.52 mm. Forefemur:

2.54 mm. Abdomen: about 2.9 mm. Head: integument of head capsule, cly-

peus, torus and flagellum brown to black; erect scales brown to black; clypeus

sometimes with a few white scales; palpus about 0.33 of proboscis; flagellar

segments 1-6 or 7 with scales. Thorax: integument of mesonotum and scutel-

lum evenly brown to black or with supraalar area somewhat lighter, postnotum

always lighter, evenly brown to beige, pleuron brown to black anteriorly, light-

er posteriorly; supraalar scale line present; lateral prescutellar and inner

dorsocentral scale lines connected; white posterior fossal scales present al-

though very few in number, and connected to outer dorsocentral scale line;

outer dorsocentral scale line with scales rather sparse; lower stp bristles 0,1

strongly developed, 4-7 weakly to moderately developed, present throughout

length of lower stp scales; part or all of mep bristles sometimes whitish in

color; pleural scaling without well defined lines; ppn scales present; psp scales

present; stp with lower vertical line of scales much broadened and extending

dorsad to upper horizontal scale row, so that entire posterior portion of stp

has sparse covering of white scales, some of which, in lower portion, may be

semierect; lower anterior mep scale patch narrow to broad, apparently never

joining upper. Legs: integument of forecoxa brown to blackish-brown, midcoxa

not or somewhat lighter, hindcoxa slightly to considerably lighter; legs. with

dark scales brown to black; forefemur with anterior surface streaked in basal

0.3-0.4, posterior surface with area dorsad of dark streak sometimes largely

dark-scaled with white speckles; midfemur with anterior surface entirely dark-

scaled with white speckles, posterior surface light-scaled in basal 0. 2-0. 4;

hindfemur with anterior surface usually light-scaled in basal 0.05-0.4; fore-

tibial anterior surface usually with line of white speckles; hindtibial ventral,

-anterior, and dorsal surfaces speckled with white scales, posterior surface

with white line in basal portion; foretarsal segment 1 usually with small dorsal

white patch at ends and white speckles basally, segment 2 sometimes with very

small dorsal white patch at ends; midtarsal segment 1 with complete narrow

basal ring and ventrally incomplete narrow apical ring, and usually with white

speckles basally, segment 2 with small dorsal white patch at each end; hind-

tarsal segment 1 usually with complete apical ring, segment 5 white-scaled on-

ly at ends or rarely only at base, patches over joints generally smaller than in

other species. Wing: light spot in area of crossveins usually poorly developed

_ Zavortink: Genus Orthopodomyia 33

and not conspicuous to unaided eye, white patches on veins R9,3, R4y,5, M and

Cu, very small or 1 or more absent. Haltere: integument of stem beige to tan,

knob brown. Abdomen: integument of tergite I beige to brown, sternites II-VII

beige to brown, sternite VIII dark brown; abdominal segments with dark scales

brown to black; tergite Il entirely white-scaled or with some dark scales apical-

ly; tergites III-VIII with basolateral white patch and sometimes, especially on

more proximal and distal segments, with a few white scales near base middor-

sally or with poorly developed basal band; sternite II entirely white- scaled;

sternites IIJ- VII with dark apicolateral patches or dark apical band, dark-

scaling usually becoming more prevalent on distal segments; sternite VIII bare.

MALE. Essentially as infemale. Labium: as in female or with white

scales forming lateral patch or complete ring just distad of level of distal end

of palpal segment 2. Palpus: about 0.93 of proboscis, segment 4 sometimes

with scattered white scales. Abdomen: as in female or with more proximal

and distal tergites with poorly developed basal bands; tergite VHI with mixed

black and white scales, sternite VIII with white apical and basal bands.

MALE GENITALIA (fig. 3). Segment VIII: tergite lobe very small, broad-

er than long, or larger, longer than broad, becoming narrower apically, apex

usually rounded, sometimes truncate with slight central emargination. Segment

IX: tergite with 1-4 bristles on each side. Sidepiece: basal mesal lobe with

3-5 stout bristles which are usually curved mesally near apex and with 1 or 2

finer bristles. Aedeagus: broader than in other species in Signifera group;

ventral triangular spines usually near middle; preapical ridge with 3 or 4 pro-

cesses. Proctiger: paraproct sclerotization usually with 2, 3 apical teeth; cer-

cal setae usually 3 or 4.

PUPA (fig. 3). Abdomen: 3.3mm. Trumpet: 0.5 mm. Paddle: 0.7 mm.

Cephalothorax: hairs 8, 9-C not as long as trumpet. Trumpet: deep tan to brown

except for extreme base and apex and largely darker than darkest part of cepha-

lothoracic integument. Metanotum: hair 12-C usually moderately developed.

Abdomen: hairs 6, 7-I medium long; 1-II moderately developed, usually 2, 3b

(1-4); 2-II-VI less than 0. 3 length of hair 1 of same segment; 6-II short and

fine, similar to 7-II; 5-III usually 2-4b (1-5) and more weakly developed than

d-IV-VI or 3-III; 10-III-VII usually not extending beyond 0.8 of following seg-

ment; 5-IV, V not extending beyond 0.5 of following segment. Paddle: usually

narrowly to broadly oval to oboval in outline; apex rounded, truncate or emar-

ginate; inner part without minute spicules near inner margin.

LARVA (fig. 4). Head: 0.96 mm. Siphon: 0.91 mm. Anal Saddle: 0.46

mm. Not all branched hairs with conspicuous barbs. Head: largely brown col-

ored, lighter cephalad and around imaginal eye, collar darker; mental plate

brown, usually with 9,10 (9-11) teeth on each side; hair 2-C apparently absent;

9-C single, fine; 11-C weakly developed, usually 3-5b; 13-C moderately devel-

oped, usually 2, 3b (2-4). Antenna: uniformly straw colored. Thorax: epider-

mal pigment moderately to strongly developed, purple in color; spicules not

conspicuous; hairs strongly pigmented; hair 0-P usually 4, 5b (3-7); 6-P sin-

gle; 1-M moderately developed, single; 1-T weakly to moderately developed,

usually single; 13-T moderately developed, usually 3b (2-4). Abdomen: seg-

ment VII usually with a large sclerotized plate and segment VI often with a

smaller dorsal plate; hairs strongly pigmented; hair 1-I weakly developed,

usually 2-4b; 4-I shorter than 3-I; 6-I usually 5, 6b; 7-I single or double; 1-II

weakly developed, usually single; 4-IJ moderately developed; 5-II-VI weakly

developed, usually single or double (1-3b); 6-II usually 5, 6b (5-8); 7-II mod-

erately developed; 1-III moderately to strongly developed, usually single; 13-

IlI-V usually single or double and either shorter or longer than hair 10 of cor-

34 Contrib. Amer. Ent. Insts; vol: 3,-no, 2, 1968

responding segment; 1-VI weakly developed, usually 1-3b; 1-VII moderately to

strongly developed, single or double; 13-VII usually 3, 4b (3-6). Segment VIII:

large sclerotized plate present; anterior comb scales usually 17-21 (14-23),

posterior comb scales usually 6-8 (6-9); posterior row of comb scales about

0.46-0. 56 length of anterior row; hair 3-VIII strongly developed, usually 8-10b

(7-11). Siphon: index about 2. 9-3, 9, usually less than 3.5; largely brown in

color, with extreme base darker and apex sometimes lighter; integumentary

sculpturing weakly developed, imbricate; hair 1-S located 0. 46-0.55 from base

of siphon, usually 7-10b (7-13), strongly developed and with longest branches

extending to or slightly beyond apex of siphon. Anal Segment: saddle complete;

saddle uniformly brown or Somewhat darker near apex dorsally; integumentary

sculpturing absent to weakly developed, imbricate; diameter of saddle at most

proximal point usually considerably smaller than diameter at most distal point

along the ventral margin, so that dorsal and ventral surfaces diverge when

viewed from the side and diameter of saddle at most proximal point usually

less than diameter of base of siphon; hair 1-X single and fine; ventral brush

usually with 7 pairs of hairs; more strongly developed hairs in ventral brush

usually with 11-14 branches (10-17); gills short to moderately long, dorsal pair

about 1.5-2.0 as long as ventral pair and 0. 45-0. 80 length of anal saddle.

BIONOMICS. Larvae of this species have been reported only from tree-

holes. Nothing is known of the habits of the adults.

SYSTEMATICS. On the basis of adult ornamentation ae is the most

derived species of the Signifera group. It appears to be a recent derivative of

signifera which arose in geographical isolation on Jamaica. It is possible that

it Should be considered as a subspecies of Signifera.

The adults are best distinguished from those of alba and signifera on the

basis of mesonotal ornamentation; in waverleyi the lateral prescutellar scale

line curves and joins the inner dorsocentral scale line and the posterior fossal

area has white scales which join with the outer dorsocentral scale line. The

adults of signifera from the Greater Antilles agree with waverleyi in having the

fifth hindtarsal segment white only at the ends and the white spot over the cross-

veins of the wing reduced, and tend towards waverleyi in ornamentation of stp.

None show the mesonotal ornamentation or other features of waverleyi. Some

signifera from Arizona and southern California also have the fifth hindtarsal

segment white only at the ends and have the wing spot reduced.

Only non-hairy forms of the larva and pupa are known. The larva is indis-

tinguishable from signifera from the eastern United States and the Greater An-

tilles and is very similar to kummi and pulchripalpis. The pupa cannot be sep-

arated from signifera from the Greater Antilles and is very similar to signifera

from California and pulchripalpis.

The mesonotum of the adult does not always show its complete ornamenta-

tion because the white posterior fossal scales and the outer dorsocentral scales

are sparse and easily rubbed off.

This species appears to be more uniform than others in the Signifera group.

This is expected as the species is endemic to one rather small island and this

island is extensively forested so that the demes are not likely to be semi-iso-

lated.

Preliminary investigation has failed to verify the past existence of a Waver-

ley Estate at Constant Spring, Jamaica, but has revealed that a Maverley Es-

tate did exist nearby. Depending upon the results of continuing research, the

name of this species may be emended to maverleyi.

DISTRIBUTION. Endemic to the island of Jamaica. Material Examined:

459 specimens; 47%, 44 2, 103 P, 265 L; 72 individual rearings (30 larval,

Zavortink: Genus Orthopodomyia 30

38 pupal, 4 incomplete).

JAMAICA. St. Andrew: Constant Spring, Waterworks gate, 7 Sept 1965,

J.N. Belkin (JA 327), 1 L [UCLA]. Constant Spring, Waverley Estate, 1 Sent

1906, M. Grabham, & lectotype, 10, 2 L [USNM]. Constant Spring, 16 Nov

1966, O.G.W. Berlin and D.C. Watson (JA 697), 21lpo (697-10,12), 1 1p?

(697-13), 1o%, 1 P, 1 L [UCLA]; same data (JA 698), 5 lpo (698-10, 12, 13, 19,

20), 6 1p? (698-11, 14, 16-18, 21), 2 po” (698-100, 102), 2 p? (698-101, 103), 2 9,

12 P; 7 {Pes}. Hermitage, 20 Aug 1964, H. L. Tucker (JA 110), we [UCLA];

21 Sept 1966, D.C. Watson (JA 625), 12 L [UCLA]. Hermitage Dam Road, 22

Aug 1965, W. A. Page (JA 280), 2 p? (280-100, 101) [UCLA]; 29 Oct 1965, Ww. A.

Page (JA 394), 1 lpo (394-10 [UCLA].

St. Mary: Annotto Bay, 16 Sept 1966, D.C. Watson (JA 622), 1 lpo (622-

10), 1 lp? (622-11), 2 po (622-100, 102), 1p? (622-101), 1 lp (622-12), 9 L

[UCLA]. Road from Annotto Bay to Broadgate, 28 Nov 1965, W.A. Page (JA

407), 5po (407-100, 102, 104,112,114), 9 p? (407-73, 74, 105-110, 112). 2,

29,6 P[UCLA]. Broadgate, Junction Road, 25 Aug 1965, W. A. Page (JA 256),

1 lpo (256-15), 1 po" (256-100), 1 p? (256-101), 1 1p (256-31), 114 L [UCLA].

Broadgate, 12 Nov 1965, W. A. Page (JA 401), 2 1lpo (401-11,13), 5 1p? (401-

14, 15, 20, 22,23), 6 po (401-100-103, 106,108), 4 p? (401-104, 105, 107, 112),

1 lp (401-21), 119 (401-12), 40%, 39,1 P, 73 L [UCLA]; 27 Oct 1966, O.G. W.

Berlin and D.C. Watson (JA 642), 3 1lpo (642-11-13), 2 1p? (642-10, 14), 2 po

(642-100, 101), 1 p¢ (642-102), 2 P, 10 L [UCLA].

St. Thomas: Leith Hall, 13 Nov 1966, O.G.W. Berlin and D.C. Watson

(JA 690), 10 P, 1 L [UCLA].

No locality or date: R.B. Hill, 1 2 [USNM].

Additional records from the literature. HAITI. Nord: Plaisance, Roche

Platte, 7-11 Oct 1925, Hoffman (Root, 1927:463,465). Since no specimens

from Hispaniola have been available for study, the identity of the species pres-

ent on the island is unknown. It is possible that the species is signifera rather

than waverleyi.

2. Orthopodomyia signifera (Coquillett)

Figs. 2,5, 6

1896. Culex signifer Coquillett, 1896:43. *TYPE: holotype 2, Washington, Dis-

trict of Columbia, United States, June, D.W. Coquillett [USNM, 3654].

1950. Orthopodomyia californica Bohart, 1950: 399-400. *TYPE: holotype 2 with

associated larval and pupal skins (48. 4,.21a), Elkhorn Ferry, near Sac-

ramento, Yolo County, California, United States, larva found in a cotton-

wood treehole, 12 Apr 1948, R.M. Bohart[USNM]. NEW SYNONYMY.

Orthopodomyia signifera of Howard, Dyar and Knab (1917:887-891); MacGre-

gor (1919:454); Edwards (1921: 290): Seguy (1921b:114); Dyar (1922:67;

1923b:96-97); Seguy (1923:155); Viosca (1923:37); Seguy (1924:201); Dyar

(1928:397-398, in part); Matheson (1929:203-204; 1930:294); Edwards (1932:

108, in part); Bishopp, Cory and Stone (1933:6); Marshall and Staley (1933:

435): Cory, Langford, Crosthwait and Graham (1934:21, 22,29); Baker

(1935:149, 151, 152,153); MacCreary and Stearns (1935: 118): Baker (1936:

1, 2, 4, 5, 7): Cae (1936:371); Clarke (1936:103); Horsfall (1936:678);

Banke (1937:146): Horsfall (1937:747); Quinby (1937:83); Shields and Miles

(1937:237); Berger (1938:834); Penn (1938:5); Pennebaker (1938:4); Shields

(1938:429); Edwards (1939:121,123); Glick (1939:51); King, Bradley and

36 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

McNeel (1939:55-56); Lane (1939:97, in part); Senevet and Abonnenc (1939:

588); Tulloch (1939:128, 130); Carpenter (1941:58); Quinby (1941:20); Reeves

(1941:69-72); King, Bradley and McNeel (1942:61); Reeves (1942:2); Rowe

(1942:501); Rozeboom (1942:43); Freeborn and Brookman (1943:22); Gurney

(1943:929, 932,935); King, Roth, Toffaleti and Middlekauff (1943:574); Knut-

son (1943:317); McGregor and Eads (1943:939); Christensen and Harmston

(1944:111); Dorer, Bickley and Nicholson (1944:49); Dorsey (1944:379, 380,

381, 382, 383, 385); Hart (1944:415); King, Bradley and McNeel (1944:61);

Matheson (1944:248-249); Middlekauff and Carpenter (1944:89); Olson and

Keegan (1944a:781, 782; 1944b:848); Pritchard and Pratt (1944:233); Quinby,

Serfling and Neel (1944:548-549); Randolph and O'Neill (1944:84); Schoof and

Ashton (1944:6, 7, 8,9); Tate and Gates (1944:24); Weathersbee (1944:643);

Good (1945:175); Headlee (1945:184-190); Kitzmiller (1945:409); Matheson

(1945:27); Peterson and Smith (1945:381); Stabler (1945:95); Carpenter and

Chamberlain (1946: 82, 83, 84, 85, 86, 87, 88); Carpenter, Middlekauff and

Chamberlain (1946:116-119); Jenkins and Carpenter (1946:40-41); Shlaifer

and Harding (1946:254); Wilson, Barnes and Fellton (1946:84); Brandenburg

and Murrill (1947:5, 6); Breland (1947a:185, 186; 1947b:81, 84, 85; 1947c:77);

Michener (1947:361); Ross (1947:36-37); Weathersbee and Arnold (1947:225);

Wirth (1947:742, 743); Brown (1948:231); Roth (1948:168); Snow (1948:87, 88);

Barnes, Fellton and Wilson (1950:82); Bohart (1950:403, 404); Fellton, Barnes

and Wilson (1950:91); Frost (1950:66); Darsie (1951:40-41); Darsie, Mac-

Creary and Stearns (1951:144); Freeborn and Bohart (1951:50); Wilkins and

Breland (1951:225, 227, 229, 233, 234); Yamaguti and LaCasse (1951:17-19);

Carpenter (1952:52); Yamaguti (1952:16,17); Hedeen (1953:3, 6); Lake (1953:

154,155); Lane (1953:628-629, in part); Masters (1953:160, 161); Sudia and

Gogel (1953:130); Venard and Mead (1953:331); Chamberlain, Sikes, Nelsen

and Sudia (1954:283); Ferguson and McNeel (1954:31); Carpenter and La

Casse (1955:101-103); Breeland (1956:101); Perez Vigueras (1956:472-474);

Pratt (1956:8); Vargas (1956:31); Breland (1957:305); Barr (1958:53-54);

Breland (1958:219, 221); Burbutis (1958:212); Chamberlain, Sudia, Burbutis

and Bogue (1958:306); Price and Abrahamsen (1958:92); Rapp (1958:28); Rut-

schky, Mooney and Vanderberg (1958:16); Breland (1959:137-141); Rapp

(1959:130); Stone, Knight and Starcke (1959:124, in part); King, Bradley,

Smith and McDuffie (1960:136-137); Vargas (1960:339, 340); Breeland, Snow

and Pickard (1961:306-307); Breland (1961:360, 372, 374); Johnson (1961:55);

Love and Goodwin (1961:214); Porter, Evans and Hughs (1961:235); Edman

(1962:431); Tinker and Stojanovich (1962:582); Dodge (1963:798, 804, 808);

Warren and Breland (1963:621, 622); Ross (1964:107, 115); Breeland and

Pickard (1965:20); Chapman (1965:435, 436, 438); Ross and Horsfall (1965:

14, 24, 43,44); Stone, Sabrosky, Wirth, Foote and Coulson (1965:109, in part);

Wiseman (1965:58); Dodge (1966:378); Flemings and Walsh (1966:426); Ger-

hardt (1966:38); Montchadsky and Garcia-Avila (1966: 36).

Bancroftia signifer of Thibault (1910:20); Seguy (1920:253).

Pneumaculex signifer of Felt (1904:490); Dyar (1905a:46; 1905b:108); Coquillett

(1906:26); Dyar (1906:3); Mitchell (1907:189-191); Theobald (1907:14, 524-

527: 1910b:469, 620).

Culex (Pneumaculex) signifer of Viereck (1908:470-471).

Mansonia signifer of Dyar and Knab (1906:185); Grabham (1907:25); Knab (1907:

153, 154). | |

Stegomyia signifera of Howard (1901:236); Theobald (1901a:322); Giles (1902:

379); Smith (1902:299); Theobald (1903:238); Aldrich (1905:126); Blanchard

Zavortink: Genus Orthopodomyia ot

(1905:258-259); Theobald (1905:19).

Aedes signifer of Grossbeck (1910:719).

Culex signifer of Howard (1896:23); Coquillett (1900:6, 7); Giles (1900:195, 268);

Howard (1900:31); Dyar (1903:26-27); Smith (1904:38; 1905:255-259); Lud-

low (1906:97). )

Orthopodomyia californica of Freeborn and Bohart (1951:50); Yamaguti and La

Casse (1951:12-17); Yamaguti (1952:16, 17); Sudia and Gogel (1953:129);

Grant (1954:73); Carpenter and LaCasse (1955:100-101); Loomis, Bohart

and Belkin (1956:42); Pratt (1956:8); Stone, Knight and Starcke (1957:123);

Loomis (1963:24); Ross (1964:107); Chapman (1965:432-439); Stone, Sabro-

sky, Wirth, Foote and Coulson (1965:108); Womeldorf and Gillies (1967:55,

56, 62).

FEMALE (fig. 2). Wing: 3.89 mm. Proboscis: 2.53 mm. Forefemur:

2.51 mm. Abdomen: about 3.2 mm. Head: integument of head capsule, cly-

peus, torus and flagellum light brown to black; erect scales dark brown to

black; palpus about 0.33 of proboscis, segments 1, 2 and 3 rarely with white

scales few and scattered (some western individuals); flagellar segments 1-4

or 0 with scales. Thorax: integument of mesonotum evenly orangish-brown,

reddish-brown or light to dark brown or fossal and/or supraalar area lighter,

scutellum evenly reddish-brown to dark brown, postnotum usually lighter, even-

ly beige to dark brown or with darker median longitudinal streak, pleuron red-—

dish-brown to dark brown anteriorly, lighter posteriorly; supraalar scale line

present or rarely completely absent (some western individuals); lateral pre-

scutellar and inner dorsocentral scale lines not connected; white posterior fos-

sal scales absent; outer dorsocentral scale line with numerous scales, rarely

curving slightly laterad at anterior end; lower stp bristles 4-10, weakly to _

moderately developed, present throughout length of lower stp scales; mep bris-

tles rarely light in color; pleural scaling with broad but usually well defined

lines; ppn scales present, or rarely reduced or absent (some western individ- »

uals); psp scales present or absent; stp horizontal scale lines usually distinct,

lower sometimes broadened posteriorly, upper and lower sometimes joined be-

fore and/or behind upper stp bristles, stp vertical scale line sometimes extend-

ing slightly dorsad of level of lower edge of mep, sometimes broadened dorsal-

ly, some upper and lower scales usually semierect; lower anterior mep scale

patch broad, sometimes continuous with upper or rarely reduced or absent

(some western individuals). Legs: integument of forecoxa light brown to brown,

midcoxa lighter, hindcoxa lightest, beige to amber; legs with dark-scaling

brown to black; forefemur with anterior surface usually streaked in basal 0. 05-

0.60, posterior surface with area dorsad of dark streak sometimes entirely

white-scaled with dark speckles; midfemur with anterior surface usually white-

scaled basally or dorsobasally, but sometimes without white scales at base,

sometimes with light scales in ventral streak from base to near middle, poste-

rior surface light-scaled in basal 0.05-0. 33; hindfemur with anterior surface

light- scaled in basal 0.05-0.4; foretibial posterior surface rarely with white-

scaling reduced to a few speckles; midtibial dorsal surface rarely speckled

with white scales (Some western individuals); hindtibial anterior, dorsal and.

posterior surfaces speckled with white scales, those on posterior surface some-

times forming a line basally; foretarsus entirely dark-scaled or with small

dorsal white patch at base of segment 1 or segments 1-2, 3 or 4 and at apex of

segment 1, or segments 1-2 or 3; midtarsal segment 1 with ventrally incom-

plete basal white ring or dorsal patch and usually with dorsal apical white patch,

segment 2 sometimes with small basal and apical dorsal white patch, segment

38 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

3 less frequently with small dorsobasal white patch, segments 4 and 5 some-

times with beige scales (some western individuals); hindtarsus with apex of

segment 1 and base of segment 2 with complete ring, apex of segment 2 and

base of segment 3 with complete or ventrally incomplete ring, segment 5 all

white dorsally or segment 5 and sometimes 4 with white-scaling much reduced,

sometimes almost absent (some western individuals and island populations).

Wing: white spot in area of crossveins well developed, conspicuous to unaided

eye, white patches on veins M, Cuy, base of R4,5 and R9,3 large or spot some-

what reduced or absent (especially in Some western individuals and island pop-

ulations). Haltere: integument of stem white to tan, knob tan to brown. Abdo-

men: integument of tergite I beige to blackish, sternites I-VII whitish to brown,

distal segments usually darker, sternite VIII light to dark brown; abdominal

segments with dark scales brown to black; tergite II with large basolateral

white patch and narrow to broad basal white band which may extend to apex of

segment middorsally; tergites III- VIII with basolateral white or dingy patch,

those on opposite sides sometimes connected by irregular dingy or distinct

white basal band; sternite II entirely white-scaled; more proximal of sternites

IlI- VII varying from entirely white-scaled to having a small to large apicolat-

eral dark patch or broad apical dark band, more distal varying from mostly

white-scaled with narrow dark apical band to mostly dark-scaled with only baso-

lateral white patches; sternite VIII with light and dark scales laterally.

MALE. Essentially as in female. Labium: as in female or with lateral or

ventral patch or narrow to broad incomplete or complete ring of white or dingy

scales just distad of level of distal end of palpal segment 2 and sometimes with

line or speckling of white scales at edge of labial groove much reduced or pres-

ent only basally or apically. Palpus: about 1.02 of proboscis. Abdomen: ter-

gites as in female, but frequency of basal banding greater, sternites generally

darker than in female, sometimes nearly all dark-scaled; tergite VIII dark-

scaled, with mixed white and dark scales, or with white apical band, sternite

VII with white basal and sometimes apical band.

MALE GENITALIA (fig. 5). Segment VIII: tergite lobe broader than long to

longer than broad, parallel sided or narrowing or widening distally, apex usu-

ally emarginate, sometimes rounded, truncate, or irregular. Segment IX: ter-

gite with 0-5 bristles on each side. Sidepiece: basal mesal lobe with 3 or 4

(2-5) stout bristles which are straight or only very slightly bent mesally to-

wards apex, and with 1-3 finer bristles. Aedeagus: ventral spines variable in

position; preapical ridge with 1-7 processes. Proctiger: paraproct scleroti-

zation with 2-5 apical teeth; cercal setae 2-5. :

PUPA (fig.5). Abdomen: 3.5mm. Trumpet: 0.4mm. Paddle: 0.6 mm.

Cephalothorax: hairs 8,9-C often as long as trumpet except in western popula-

tions. Trumpet: deep tan to brown except for extreme base and apex and large-

ly darker than darkest part of cephalothoracic integument. Metanotum: hair

12-C often strongly developed. Abdomen: hairs 6, 7-I usually strongly devel-

oped and long in all but most Californian and some Puerto Rican individuals;

1-II usually moderately developed and 4, 5b (4-16) in eastern populations, often

strongly developed, long and 2, 3b (1-5) in western populations; 2-II-VI not

more than 0.3 length of hair 1 of corresponding segment; 6-II moderately to

strongly developed and stronger than 7-II in all but most Californian and some

Puerto Rican individuals; 5-III usually 2, 3b (1-5) and not as strongly developed

as 5-IV-VI or 3-III; 10-III, VI sometimes extending to apex of following seg- —

ment, 10-IV,V, VII usually not extending beyond 0.8 of following segment; 5-IV,

V extending to about 0.4 to 0.6 of following segment in eastern populations but

extending to 0.7-1.0 of following segment in western populations. Paddle: usu-

Zavortink: Genus Orthopodomyia og

ally narrowly elliptic, oval or oboval in outline in eastern populations, usually

broadly oboval in outline in western populations; apex rounded, truncate or

emarginate; inner part without band of minute spicules near inner margin.

LARVA (fig.6). Head: 0.95 mm. Siphon: 0.75 mm. Anal Saddle: 0.47 mm.

Not all branched hairs with conspicuous barbs. Head: largely light brown to

brown, lighter cephalad and around imaginal eye, collar darker; mental plate

brown, usually with 8-10 teeth on each side; hair 2-C apparently absent; 9-C

usually single (1-3b), fine; 11-C weakly developed, usually with 3, 4b (2-6);

13-C moderately developed, usually 2, 3b (1-4). Antenna: uniformly light tan

colored or lighter towards apex or base. Thorax: epidermal pigment usually

strongly developed, purple, in eastern populations, but absent or weakly devel-

oped, pink or reddish, in western populations; spicules not conspicuous except

in hairy forms; hairs strongly pigmented; hair 0-P usually 3-5b (2-6); 6-P sin-

gle; 1-M usually moderately developed and single ineastern populations, usual-

ly strongly developed and single or double (1-4b) in western populations; 1-T

usually very weakly developed and single or double in eastern populations, mod-

erately to strongly developed and usually single or double (1-4b) in western pop-

ulations; 13-T usually moderately developed and 2, 3b (2-4) in eastern popula-

tions, strongly developed and usually 2, 3b (2-5) in western populations. Abdo- —

men: segment VII usually and segment VI often with sclerotized plate; hairs

strongly pigmented; hair 1-I usually very weakly developed and 2, 3b (2-4) in

eastern populations, usually moderately to strongly developed and 2-4b (2-6)

in western populations; 4-I shorter than 3-I; 6-I usually 4-7b (4-8); 7-I usually

single or double in eastern populations, usually double or 3b (1-5) in western

populations; 1-II usually very weakly developed and single or double (1-3b) in

eastern populations, usually moderately to strongly developed, single or double

(1-3b) in western populations; 4-II usually moderately developed, but strongly

developed in hairy forms; 5-II-VI usually weakly developed and single or double

in eastern populations, usually moderately to strongly developed and 1-3b (1-4)

in western populations; 6-II usually 5, 6b (2-8); 7-II moderately developed; 1-

III usually moderately developed and single (single, double) in eastern popula-

tions, usually moderately to strongly developed and single or double (1-3b) in

western populations; 13-III-V usually single or double (1-3b) and shorter than

hair 10 of corresponding segment in eastern populations, usually 1-3b (1-5) and

longer or shorter than hair 10 of corresponding segment in western populations;

1-VI usually weakly developed and 2, 3b (2-4) in eastern populations, usually

moderately to strongly developed and 2, 3b (1-4) in western populations; 1-VII

moderately to strongly developed, usually 2, 3b (1-4); 13-VII usually 2-4b (2-6).

Segment VIII: large sclerotized plate usually present; anterior comb scales

usually 17-21 (11-24), posterior comb scales usually 6-8 (5-11); posterior row

of comb scales about 0. 40-0. 75 length of anterior row; hair 3-VIII strongly de-

veloped, usually 5-7b (3-8). Siphon: index about 2.3-3.8, usually 2.7-3.5;

largely brown in color, extreme base somewhat darker, sometimes area basad

of hair 1-S slightly lighter, apex not or somewhat lighter; integumentary sculp-

turing absent or weakly developed, imbricate; hair 1-S strongly developed, lo-

cated 0. 36-0.48 from base of siphon, usually 8, 9b (6-10) and with longest

branches extending almost to apex of siphon in eastern populations, usually 6,

Tb (5-10) and with longest branches extending far beyond apex of siphon in west-

ern populations, and 9-13b and with longest branches extending far beyond apex

of siphon in Cuban specimens. Anal Segment: saddle complete; saddle brown,

somewhat darker dorsoapically; integumentary sculpturing weakly developed,

imbricate; diameter of saddle at most proximal point slightly to considerably

smaller than diameter at the most distal point along the ventral margin, so that

40 | | Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

dorsal and ventral surfaces diverge when viewed from the side and diameter of

saddle at most proximal point equal to or less than diameter of base of siphon;

hair 1-X usually single (1-3b), fine; ventral brush usually with 7 pairs of hairs;

more strongly developed hairs in ventral brush usually with 8-12 branches (6-

13); gills usually moderately long, dorsal pair about 1.5-2.4 as long as ventral

pair and 0.6-1.2 (0.6-1.5) length of anal saddle.

BIONOMICS. Larvae of this species have been found in rot holes in trees

and stumps, in artificial containers, particularly wooden ones or those con-

taining twigs and leaves, and in flooded cesspools and manure pits. They are

frequently found in association with larvae of alba where the ranges of the two

Species overlap and can as found in the same treeholes as kummi in southeast-

ern Arizona.

Adults have been caught in light and malaise traps and have been collected

as they rested inside rot holes or on the bark of trees. Glick (1939:51) reported

catching 1 adult at 500 feet altitude in a trap on the wing of an airplane.

Thibault (1910:20) reported that females entered dwellings and bit frequent-

ly in Arkansas. This obServation has never been verified by others. Females

are known to take avian blood. Vargas (1960:339, 340) found adults from Tamau-

lipas, Mexico, infected with the virus of Eastern Equine Encephalitis.

For additional information on the biology and ecology of western populations

of this species see Chapman (1965:432-439) and the Biology and Ecology section

of this paper. 7

SYSTEMATICS. This is the most widespread and variable species in the

Signifera group. The treatment given here differs from that of other authors

in that I am considering californica Bohart to be a synonym of signifera and

waverleyi Grabham to be a distinct species.

Adults of signifera are quite easily told from those of waverleyi by their

mesonotal ornamentation. They are likely to be confused only with those of al-

ba, but can be told from them by the presence of 4-10 bristles among the lower

stp scales. In addition, in the eastern United States, where this species is

sympatric with alba, vein R4,5 is usually white-scaled at the base whereas it

is normally dark-scaled in alba.

Non-hairy, semi-hairy (=californica), and hairy forms of the larva exist.

Non-hairy larvae are impossible to separate from waverleyi and are difficult

to separate from kummi and pulchripalpis. Semi-hairy individuals are easily

separated from the larvae of all other species in the Signifera group but the

hairy form is similar to that of kummi. Both hairy and non-hairy forms of the

pupa are known, the former very similar to the normal pupa of —S and the

latter very similar to those of waverleyi and pulchripalpis.

The range of signifera can be divided into 4 major areas, each of which is

characterized by a more or less unique combination of larval and pupal forms

and has a different pattern of adult variation. Throughout the eastern United

States and as far west as Texas and northeastern Mexico adults, pupae and lar-

vae are fairly uniform, with a large proportion of the total variation for this

broad geographical area being found within any one deme. Larvae from this

area are almost always deeply pigmented and non-hairy, whereas pupae are

nearly always hairy and have narrow paddles. Adults are characterized by

having the white rings over the joints between hindtarsal segments 1-2 and 2-3

covering a greater portion of the apex of the proximal segment than of the base

of the distal segment.

The second area lies to the west and includes New Mexico, Arizona, Utah

and the portion of California along the Colorado River. Populations in this arid

area are often quite isolated, with the result that they differ considerably from

Zavortink: Genus Orthopodomyia : 41

each other. The larvae throughout this area are not or only slightly pigmented

and are usually semi-hairy, although both hairy and non-hairy larvae do occur.

Pupae are usually hairy, as in the eastern United States, but have some hairs,

especially 1-II and 5-IV,.V, often nearly as strongly developed as in kummi.

The pupal paddles are wider distally and more emarginate apically than in sig-

nifera from the eastern United States. The most numerous and striking adult

variations of the species are found in thisarea. Banding of hindtarsal segments

1-2 and 2-3 may be as in eastern signifera, or as in Californian signifera (see

below) or intermediate. Hindtarsal segment 5 may be white only at the ends or

at the base and segment 4 often has its white-scaling reduced. The white spot

over the crossveins of the wing is more frequently reduced or absent than in

the other areas. The most conspicuous variation involves the supraalar, ppn

and lower mep scales, which may be completely absent in some individuals

from central and southeastern Arizona.

The third area is California west of the Colorado River drainage and possi-

bly Oregon. Here, as in the eastern United States, all stages are quite uniform,

probably indicating that populations are not isolated. The larvae from this area

are not or only lightly pigmented and are Ssemi-hairy; they are very similar to

those from the New Mexico-Arizona area. The pupae are normally non-hairy,

unlike those of either the first or second area, but have the paddles broadened

distally as in those from the New Mexico-Arizona area. Normal adults are

identical to signifera from the eastern United States except that the rings over

hindtarsal joints 1-2 and 2-3 often cover equal portions of each segment in-

volved. Some specimens from San Diego County, California, have the last hind-

tarsal segment white-scaled only at the ends, but this is probably due to the de-

pauperate nature of the individuals. The Californian signifera is what Bohart

described and named as californica in 1950 on the basis of larval characters.

The fourth area to be considered is the Greater Antilles. Signifera is ap-

parently present on all the larger islands except Jamaica, which is inhabited

by waverleyi. Although very few specimens from this area are available for

study, it is evident that the populations on the different islands very consider-

ably from each other. However, the larvae from all islands agree in being

highly pigmented and non-hairy, as in the larvae from the eastern United States.

The pupae are non-hairy to slightly hairy and have narrow paddles; they resem-

ble the Californian population in chaetotaxy but resemble eastern signifera in

paddle shape. Larvae from Cuba have shortened siphons, more strongly devel-

oped hair 1-S, and longer gills; in the latter 2 characters they resemble kummi.

Adults throughout the area seem to always have the last hindtarsal segment

white only at the ends, often have the spot over the crossveins of the wing re-

duced or absent and have the scales on stp more scattered. In all of these re-

spects they resemble waverleyi, but none show the mesonotal ornamentation or

other characteristics of that species. |

As discussed in the systematics section for the Signifera group, I am not

considering the various larval and pupal forms from the 4 different geographi-

cal areas to be distinct species. Most of the larval and pupal differences are

restricted to chaetotaxy and the differences are of the same type as those found

between hairy and non-hairy forms of several other species of Orthopodomyia.

The pattern differs in signifera, however, in that the hairy pupal and semi-

hairy larval forms are apparently genetically fixed over a portion of the range

and are largely allopatric from the non-hairy ones. All of the adults except

those from the Greater Antilles and the form lacking supraalar scales from

Arizona are very Similar and even these variants are more similar to typical

signifera than are the adults of any other species in the Signifera group except

alba. :

42 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

The general pattern of variation in larval and pupal chaetotaxy can be ex-

plained if one assumes that at the timeof its spread throughout the United States

and the Greater Antilles signifera had only non-hairy larvae and pupae. Later

a semi-hairy larval form became fixed in the western United States and a hairy

pupal form arose in the eastern United States or was obtained by introgression |

with kummi. The semi-hairy larval form has not spread farther eastward than

New Mexico and the hairy pupal form has not reached the populations in Calif-

ornia or the Greater Antilles.

Ornamentation of adults from California and Arizona, and presumably from

the rest of the range, varies with larval nutrition. Poorly fed larvae give rise

to depauperate adults which have the white-scaling on all parts of the body re-

duced and abundantly fed larvae produce adults with superabundant white scales.

As mentioned above the most conspicuous adult variation is found incentral

and southeastern Arizona where individuals lacking the supraalar, ppn and low-

er mep scales may be found. In all cases this form has been found in the same

treehole as normal signifera. A few individuals lacking supraalar scales have

the ppn and the lower mep scale patches reduced rather than completely absent

and some adults with supraalar scales have the patch on ppn reduced. The lar-

va of the form lacking scales is identical to that of sympatric typical signifera

and shows the same pattern of geographical variation in chaetotaxy as does the

latter. The pupa usually has hairs 5-IV, V slightly longer than in typical signi-

fera from the same treehole. The greatest number of specimens of this form

have been reared from larvae collected at 2 different times (March and Septem-

ber, 1966) from one treehole in Cochise County, Arizona. Five out of 54 (9 %)

of the mass reared pinned adults from the first collection and 22 out of 223

(10 9%) of those from the second collection lacked the scales in question. I am

considering this form to be a variant of typical signifera rather than a distinct

species.

Hybrids between signifera and kummi have been found in Arizona and are

discussed more fully under kummi. While the geographical variation of wing

scaling in kummi can be interpreted as indicating introgression of signifera

characters, I have seen no evidence of introgression of kummi characters into

signifera. However, the type of development of pupal hairs 1-II and 5-IV, V in

signifera from the Arizona, New Mexico, southeastern California and Utah area

and possibly even the typical hairy pupal form of signifera may have been ob-

tained from kummi during past introgression.

DISTRIBUTION. Oregon, Utah, New Mexico, North Dakota and Massa-

chusetts, south to California, Tamaulipas, Cuba and the Virgin Islands. Ma-

terial Bxamined: 7508 specimens; 1203 0, 987 9, 1 A, 1 gynandromorph, 6—

isolated & genitalia, 1968 P, 3392 L; 560 individual rearings (438 larval, 36

pupal, 86 incomplete).

CUBA. La Habana: Marianao, 1936, H. P. Carr, 1, 1 9, 1 gynandromorph,

4 L [USNM].

No locality: 21 June 1939, H. P. Capp, 10, 19 [USNM ].

MEXICO. Coahuila: Allende, 2 Sept 1960, "M. Yebra, 2 L [ISET].

PUERTO RICO. Humacao: El Yunque, 24 Aug 1942, GE. Bohart;, 6 ¢;

32 [CAS]; 30 Aug 1942, G.E. Bohart and T.H.G. Aitken, 1 P, 2 L [UCLA];

8 Sept 1942, 2 L [UCD], 1 L [CAS]; 21 Oct 1945, H.D. Pratt, 1 L [USNM]; 12

Jan 1946, J. Rozas, 1 P [USNM], 1 L [CU].

UNITED STATES. Alabama: Cave Springs, Wheeler Lake, 10 June 1942,

J,N. Belkin, 1 lpo (164); 2:1p? (97,99), 1 po.(141), TPlce (139), Llp (165),

fo JUCLA};8 July 1942, JUN. Belkin, Isp? (178) 1¢, 19, 1 P [UCLA].

Ozark, 2 Apr 1943, 2 L [UCLA], 2 L[USNM]. Ozark, Camp Rucker, 26 Feb-

Zavortink: Genus Orthopodomyia } 43

Apr 1943, J.G. Franclemont, 160%, 79 [CU]. Point Clear, 9 May 1953, W.L.

Seal, 3L [USNM]. Sheffield, 18 Aug 1942, J.N. Belkin, 3 p? (355, 356, 357),

20 [UCLA]; 27 Aug 1942, J.N. Belkin, 3 o [UCLA]. Wilson Dam, 14 Apr 1937,

6% [CU]; Aug 1953, R.X. Schick, 2 L [UCLA]; 21 Aug 1954, R.X. Schick, 1¢,

29 [UCLA].

Arizona: Camp Verde, 1 Sept 1966, T.J. Zavortink (UCLA $16), 2 ipg

(316-10, 11), 4 1p? (316-12-15) [UCLA]; same data (UCLA 317), 21po (317-11,

13), 41p2 (317-12, 14-16), 2 po’ (317-102, 104), 3 p9 (317-100, 101, 103), 1 Ip

(31%-10),.2 P [UCLA]. Chiricahua National Monument, 24 Dec 1966, L. T.

Nielsen (N-36-66), 1 o [UTAH]; same data (N-37-66), 3 0", 29, 1A, 6L [UTAH];

same data (N-38-66), 30 o, 119, 2 P, 14 L[UTAH]. Globe, 22 Mar 1966, L. T.

Nielsen (N-9-66), 30°, 13 L [UTAH]. Globe, Jones Water Campground, 2 Sept

1966, T.J. Zavortink (UCLA 331), 3lpo (331-10, 14,15), 12 1p9 (331-11-13,

16-24), 41%, 2392, 70P, 92L [UCLA]. Mammoth, Aravaipa Canyon, 21 Dec

1966, L. T. Nielsen (N-35-66), 30°, 4 P, 3 L [UTAH]. Pearce, Cochise Strong-

hold Campground, 26 Aug 1964, J. Burger (UCLA 255), 7 po (255-102, 104, 107-

111), 4 p@ (255-100, 101, 105, 106), 70, 29 [UCLA]; 22 Mar 1966, T.J. Zavor-

tink (UCLA 302), 1 po (302-17), 1¢, 29, 3P, 3L [UCLA]; 4 Sept 1966, T.J.

Zavortink (UCLA 326), 2 lp (326- 38, 39), 1-G [UCLA]; same data (UCLA 328),

21po (328-29, 30), 13 lp? (328-11- 14, ie 18-20, 23, 24, 26-28), 51p (328-10,

15,47 1-21) 25): 3 S19) de? 2 [UCLA]; same data (UCLA 329), 6 Ip? (329-

14- 19), 7 o1e [UCLA]; 6 Saye 1966, T.J. Zavortink (UCLA 342), 2 lpo (342-

24,28), 4 1p9 (342-17, 22, 23, 25), 1c, 1 P, 1 L [UCLA]. Phoenix, Verde Riv-

er-Salt River Recreation Area, 1 Sept 1966, T.J. Zavortink (UCLA 318), 31p?

(318-10-12) [UCLA]; same data (UCLA 319), 1 lpo (319-11), 1 lp (319-10), 1 P

[UCLA]. Portal, SW Research Station, 5-25 Sept 1965, C.W. Sabrosky, 1 o

[USNM]. Portal, 6 Sept 1966, T.J. Zavortink (UCLA 343), 1 lpo" (343-25), 19

Ip? (343-11, 18, 20-24, 26-28, 50, 52-59), 3 lp (343-10, 29, 51), 42 0, 249, 73 P,

90 L [UCLA]; same data (UCLA 344), 5lpo& (344-11, 12, 15, 16, 21), 6 1p? (344-

13,14,17-20), 50°, 5P, 8L[UCLA]. Prescott, 1 Sept 1966, T.J. Zavortink

(UCLA 315), 6 lpo” (315-17, 19-21, 23, 24), 3 1p?°(315-12-14), 1 p? (315-100),

5 lp (315-10, 11, 15, 22, 25), 2, 39, 9 P, 9 L[UCLA]. Rimrock, 20 Mar 1966,

L. T. Nielsen (N-6-66), 30°, 19, 2 P, 7L [UTAH]. Rimrock, Beaver Creek

Campground, 20 Mar 1966, L.T. Nielsen (N-7-66), 20, 22, 10 L [UTAH].

Sedona, 19 Mar 1966, L. T. Nielsen (N-3-66), 20°, 39, 9 L[UTAH]. Sedona,

Chavez Crossing Campground, 20 Mar 1966, L. T. Nielsen (N-4-66), 20, 2L

[UTAH]. Show Low, 14 Aug 1966, L. T. Nielsen (N-32-66), 1 9 [UTAH]. Sierra

Vista, San Pedro River, 22 Mar 1966, T.J. -Zavortink nec 301). 27 Ips

(301-10, 22, 25, 26, 36, 41, 45, 50, 51, 53-55, 57, 60, 61, 63-65, 68, 71-75, 77, 79, 83),

33 1p (301-11, 13, 16-21, 27, 30, 37-40, 43, 44, 46, 47, 49, 56, 58, 59, 67, 69, 70, 76,

78, 81, 82, 84-87), 18 lp (301-12, 14, 15, 23, 24, 28, 29, 31-35, 42, 48, 52, 62, 66, 80),

54 o', 329, 136 P, 136 L [UCLA]; 5 Sept 1966, T.J. Zavortink (UCLA 336),

2 1po (336-35, 39), 21 1p? (336-12-16, 19, 21-29, 32-34, 36-38), 7 1p (336-10, 11,

17, 18, 20, 30, 31), 108 o, 692, 210 P, 156 L [UCLA]; same data (UCLA 337),

1 lpo (337-11), 5 1p (3837-12-16), 70, 69, 19 P, 21 L [UCLA]. Wickenburg,

29 Dec 1965, L. T. Nielsen (N-36-65), 11 0, 89, 6 P, 8 L [UTAH]; 18 Mar 1966,

Eee, wicket (N-1-66), 20, 49, 13 L [UTAH]; 19 Dec 1966, L. T. Nielsen

(N-34-66), 3 o& [UTAH].

Arkansas: Fort Smith, Camp Chaffee, 21 Sept 1943, B. Crafchick, 10

[CU]. Little Rock, Boyle Park, 28 Mar 1943, J.N. Belkin, 1 P [UCLA]. Little

Rock, Little Fourche Bayou, 29 Mar 1943, J.N. Belkin, 2 P [UCLA]. Pine

Bluff, 26 Mar 1925, C.W. Fant, 2? [USNM], 12 [BMNH]. Scott, 24 Aug 1909,

J.K. Thibault, 30, 1 9 [USNM].

AA Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

California: Bakersfield, 17 May 1948, E. Hill, 6 L [UCD]; 2 May 1954,

w.A. McDonald (UCLA 104), 41po’ (104-113, 119, 302, 305), 14 1p? (104-102,

104-108, 111, 112, 116-118, 120, 301, 303), 31p (104-103, 109, 306), 18 c, 15 Q,

64 P, 41 L[UCLA], 1 1po' (104-110), 2 1p? (104-114, 115) [USNM]; 21 Oct 1954,

W.A. McDonald and J. Chao (UCLA 148), 110, 59, 30 P, 75 L [UCLA]. Be-

necia, C. Ludlow, 1 o [BMNH]. Calabasas, Tapia Park, 19 Oct 1963, T.J.

Zavortink (UCLA 417), 21lpo (417-20, 22), 41p9 (417-23-26), 1 1p (417-21),

30, 3P, 5L[UCLA]. Elkhorn Ferry, Yolo Co., 16 Feb 1948, R.M. Bohart,

12 L [UCD], 2 L [USNM]; 12 Apr 1948, R.M. Bohart, 12 lpo (48.4. 26a, b,c, e,

oh, 48.4.27a,c,d, 48.5.6a,c,f), 31p9 (48.4.27b, 48.5. 6d, e), 2 1p (48. 4. 26d,

f), 40°, 1 L [UCD], lp holotype of californica (48.4.21a), 1 lpo, 2 1p? (48.

4.21c,d, 48.4. 22b), 1 o, 3 L [USNM]; Apr 1948, R.M. Bohart, 4 9 [USNM], 1¢,

1 9 [BMNH]; 12 May 1948, R.M. Bohart, 2 L [BMNH]; 5 Aug 1948, R. M. Bohart,

1 & [UCD]; 25-26 Aug 1948, R.M. Bohart, 1, 6 9 [UCD], 20", 29, 2 L[USNM],

1c’, 12 [BMNH]; 3 Nov 1949, R.M. Bohart, 3L [UCD], 3L [USNM]; 3 Nov

1950, R.M. Bohart, 10 “, 49 [UCD]; 4 June 1965, T.J. Zavortink (UCLA 421),

1 1p (421-10), 1 p? (421-100), 2 P, 6 L [UCLA]; same data (UCLA 422), 39

[UCLA]. Helendale, 8 Jan 1966, T.J. Zavortink (UCLA 425), 10 lpo (425-10,

11, 18-20, 22, 23, 26-28), 8 1pQ (425-14-17, 21, 24, 25, 29), 2 1p (425-12, 13), 18 ¢,

99, 35 P, 91L [UCLA]. Kern Co., 12 Apr 1949, D. Marquez, 2 L [HARM].

Needles, 19 Feb 1966, T.J. Zavortink (UCLA 294), 9 lpo’ (294-10-13, 16, 17, 20,

21, 26), 7 1p? (294-18, 19, 22-25, 28), 3 lp (294-14, 15, 27), 9%, 102, 19 P, 30 L

[UCLA]; same data (UCLA 295), 2 1po (295-12, 13), 2 1p (295-10, 11), 1 L[UCLA];:

26 Mar 1966, T.J. Zavortink (UCLA 307), 2lpo’ (307-11, 12), 31p? (307-10,

13,14), 1¢, 1 P, 2L [UCLA]; same data (UCLA 308), 1 po’ (308-100), 31p

(308-10-12), 3 L [UCLA]; 29 Dec 1966, T.J. Zavortink (UCLA 414), 5 po

(414-12-14, 17,18), 4 1p (414-10, 11, 15,19), 1 lp (414-16), 110, 22, 14P,

76 L [UCLA]. Nice, Lake Co., 22 Feb 1942, A.W. Lindquist, 2 o [USNM].

Olancha, 13 Apr 1966, T.J. hawwetink (UCLA 309), 1 1p (309-10), 1 L [UCLA].

Pacheco, Contra Costa Co., 16 June 1954, J. Mallars, 1 L ype Pala, 3

Feb 1966, T.J. Zavortink (UCLA 293), 12 ipo (293-20- 27, 29,50, 52,55), Lip?

(293-54), 31p (293-28, 51,53), 10, 3P, 19L [UCLA]. Dearblossom: 8 Jan

1966, T. J. Zavortink (UCLA 424), 1 ow, 10 9, 14 P, 7L [UCLA]. Piedra, Fres-

no Co., ‘T Mar 1962, H.C. Chapman, 6 Bio B® [UTAH]. Pine Flat Dam, Fres-

no Co., 15 Aug 1966, Kleiwer and Clark (569), 19 [UTAH]. Piru, Lake Piru, 9

Feb 1964, T.J- Zavortink (UCLA 418), 11 ¢, 39, 55 L [UCLA]; 17 Oct 1964,

T. J. Zavortink (UCLA 419), 6 lpo’ (419-11, 13, 14, 16-18), 4 1p (419-10, 12, 15,

20), 1 1p (419-19), 10, 49, 8P, 234L [UCLA]; 21 May 1965, T.J. Zavor-

tink (UCLA 420), 31po (420-10, 11,13), 5 lp? (420-12, 15-17,19), 2 1p (420-

14,18) [UCLA]. Ravenna, Soledad Campground, 11 Dec 1965, T.J. Zavortink

(UCLA 423), 6 lpo (423-10-12, 14, 15,17), 5 1p? (423-13, 16, 18-20), 14.0, 2,

17 P, 13 L [UCLA]; 23 July 1966, T.J. Zavortink (UCLA 314), 1 1po (314-10),

23%, 119, 51 P[UCLA]. Redlands, 18 July 1940, 19 [CU]. Riverside, 2 Mar

1940, W.C. Reeves, 1 o [UTAH]; 3 Apr 1940, W.C. Reeves, 1 2 [UTAH]; 13-

21 June 1940, W.C. Reeves, 1 0, 1 9 [BMNH], 1 o, 1 ¢ [CAS], 1 o, 1 2 [USNM],

1% [CU]; 10 July 1940, W.C. Reeves, 1¢ [CAS]; 1 Sept 1944, W.C. Reeves,

20°, 29 [UTAH]; 17 May 1948, W.C. Reeves, 2 L [UCD]; W.C. Reeves, 50

[USNM]. Riverview Park, Fresno Co., B. Rosay, 2 L [USNM].

Delaware: Bombay Hook, 4 Sept 1963, R. W. Lake, 2 0, 3 9 [UTAH]. Dela-

ware City, 22 June 1933, D. MacCreary, 10 [USNM]; 5 Sept 1934, D. Mac

Creary, 1 “ [USNM]. Lewes, 1 Aug 1935, 1 &o [USNM]. Millsboro, 31 Aug 1935,

D. MacCreary, 1 2? [USNM].

District of Columbia: 12-20 Oct 1903, T. Pergande, 40, 2 2 [USNM];

Zavortink: Genus Orthopodomyia 45

June-July 1904, H.G. Dyar, 1 P, 2 L [USNM]; 26 Sept 1942, 1 o [USNM]; 22

Oct-7 Nov 1942, 1 o& [USNM]; 27 May 1943, L. T. Gallinger, 1 o [USNM]; 3

Aug 1943, 19 [USNM]; 16 Oct 1943, J. Kenilworth, 1% [USNM]; 25 May, HLS.

Barber, 1 @ [USNM]; June, D. Coquillett, ? holotype of signifera [USNM]; 26

Aug, H.S. Barber, 1 o [USNM]; Sept, W.V. Warner, 2, 2 9 [USNM].

Florida: Benson, 8 Apr 1943, S. F. Bailey, io genitalia [UCD]. Orange

Co., 1 May 1937 (1862), 1o%, 19 [USNM]. Pensacola, 8 Apr 1943, Bronson,

2 tk [UCD]. Rock Springs, Orange Co., 28 June 1937 (1883), 1192 (1883-1),

1c’, 2 9, 3L [USNM]. No locality, 1 Nov 1935, 219 (1708, 1709-1) [USNM].

No locality or date (1664), 1 2 [USNM].

Georgia: Atlanta, 9 July 1943, J. B. Duncan, 2 & [UTAH]; 30 Jan 1944, 1<¢,

2° [USNM]; 23 May 1944, 1L [USNM]: May 1944, 1, 192 [USNM]; Sept 1946,

1 L [UCLA]; 14 Jan 1947, 19 [CNC]; 13 Aug 1947, W.D. Sudia, 1 L [UTAH]:

Oct 1947, H.D. Pratt, 2o', 19 [CNC]; 13 Aug 1949, H.D. Pratt, 4 L [UTAH].

W.B. Summerall, 10, 12 [USNM]. Atlanta, Fort McPherson, 23 Mar-8 Apr

1944, 40°, 6? [USNM]; 30 Apr 1944, 2 L [USNM]; May 1944, 1 L [USNM]. Co-

lumbus, Fort Benning, 1 2 [USNM]. Newton, Mar 1949, 30°, 39 [UCLA]. Sa-

vannah, 8 Feb 1932, 2 L [USNM]; 20 Feb 1933 (40), 1 o& [USNM]; 7 Mar 1933,

Io [USNM]; 16-30 Oct 1933, 1, 59 [USNM]; 15-20 Oct, V.H. Bassett, 1 ?

[USNM]. Spring Creek, Decatur Co., 16-29 July 1912, 1% [CU]. No locality,

1956, A.R. Barr (244), 2L [USNM]. No locality or date, 5 L [HARM].

Illinois: Glencoe, July 1925, C.W. Edwards, 6 & [USNM].

Indiana: No locality, July 1942, 20, 29 [cu]. No locality or date, 2 L

[HARM], 1 o genitalia [UTAH]. |

Iowa: Ames, 28 Aug 1919, E.W. Laake, 1 2 [USNM]; 1 Aug 1947, J. Laf-

foon, 3 1p (1115. 3, 1115. 4, 1115. 5) [UCD].

Kansas: Oberlin, 19 Sept 1963, F.C. Harmston, 50, 692 [UTAH]; 9 L

[HARM]. —

Louisiana: Alexandria, 26 Oct 1942, J. Zukel, 1% [USNM]. Baton Rouge,

5 Apr-12 May 1903 (94), 29 [USNM]; 15 Aug 1941, W.W. Wirth, 4 L [USNM].

Buras, 12-13 Mar, W.V. King, 1 2? [USNM]. Kilbourne, 15 May 1944, W.W.

Wirth, io,73 2 [USNM]. Lake Charles, 15 June 1943, W. W. Wirth, 2 mist <

[USNM]. Leesville, Camp Polk, May 1945, K.L. Maehler, 2 &, 32 [CAS].

Mound, 26 Aug-2 Noy 1914, Hintey (4340), 1%, 59 [USNM]; Sept 1929, 19

[USNM]; (9676), 19 [USNM]; (9682), 2 o [USNM]; (10180A5), 1 9 [USNM]. New

Orleans, 2 Jan 1913, W.V. King (142), 1 S[USNM]. New Orleans, Camp Plauche,

Apr 1944, 3 9 [USNM]. Norco, 15 Oct 1944, F.N. Young (208- 9), 12 L, 2P

[UCLA], 2 L [CAS]. Pineville, 2 Nov 1942, T.S. Ford, 1 o [USNM]. Simms,

Camp Livingston, 1 Aug 1942, W. Ww. Fassig, 21L [CAS].

Maryland: Annapolis, 15 ‘Aug 1932, 19 [USNM]; 27 July 1933, F.C. Bish-

opp, 12 [USNM]. Baltimore, Patapsco State Park, 17 Oct 1965, W.A. Mc

Donald (UCLA 286), 9 lp&% (286- 03, 55, 62, 64, 65, 70, 72. $1,.82),.13 1p? (286-52,

04, 57-59, 61, 63, 68, 69, 74, 78, 84, 85), 4 po (286- 206,11; 73, 76), 3 pe (286-56, 67,

75), 6 1p (286- 50, 51, 60, 77, 80, 83), LT. Bae 73 P, 91 L [UCLA]. Bethes-

da, 29 July 1944, G. B. Vost, 4 sie [USNM]; a2 Aug 1944, G.B. Vogt, 1¢%

[USNM]. Cabin John: 5-15 Sept 1908, F. Knab, 5 o [USNM]; Oct 1903, F. Knab,

20°, 19 [USNM]; 7- 13 Sept, F. Knab, 2 0 [USNM]. College Park, 17 July 1932,

F. C. Bishopp, 1 2? [USNM]. Piantiiers Island, 7 June 1906, D.H. Clemons,

12 [USNM]. Riverview, Aug 1905, T. Pergande (9910), 7 0’ [USNM].

Massachusetts: Boston, Fort Strong, 5 Oct 1924, 72 [USNM].

Mississippi: Bay Saint Louis, 8 June 1913, W. V. King (142), 4 0, 2 97USNM].

Electric Mills, J.A. Le Prince, 3 2 [USNM]. Hattiesbure, Camp Shelby, 27 Mar

1944, C.D. Michener, Bil; [USNM]. Perkinston, 14 Sept 1933, 1 9 [USNM].

46 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

Missouri: Eagle Rock, Roaring River State Park, 5 July 1942, A.B. Gur-

ney, 3 L[USNM]. Neosho, Camp Crowder, 3 July-30 Aug 1942, A.B. Gurney,

4% [USNM]. St. Louis, Aug-Oct 1904, A. Busck, 15, 10 2? [USNM].

Nebraska: Niobrara, 1 L [HARM].

New Jersey: Chester, 8-10 Sept, 22 [CU]; 10-13 Nov, J.M. Aldrich, 39

[USNM]. New Brunswick, J.B. Smith, 1%, 1 29 [USNM].

New Mexico: Bernalillo, 19 Mar 1967, L. T. Nielsen and J.H. Linam (N-

2-67), 3o%, 2 P, 3L [UTAH]. Glenwood, Catwalk Campground, 21 Mar 1967,

Lot: Nieleert nnd H. Linam (N-5-67), 4 a 52, 10P, 7 L[UTAH]. Los Lanes

20.Mar 1967, L. T. Nielsen and J. H. Linam (N- 3-67), 4 &,1°92,6P,4-L [UTAH].

Reserve, 21 Mar 1967, L. T. Nielsen and J.H. Linam (N-4b-67), 4 o, SBP,

8 L [UTAH]. Socorro, Escondida Recreation Area, 2 Sept 1966, T.J. Zavor-

tink (UCLA 320), 6 lpo& (320-10-12, 15,17, 18), 41p9 (320-13, 14, 16,19), 41 ¢,

16 9, 61 P, 156 L [UCLA]; same data (UCLA 321), 7 1lpo (321-10, 11, 14-16, 18,

19), 3 1p (321-12, 13,17), 42 0, 239, 69P, 157 L [UCLA]; same data (UCLA

329). Tl po (322-16): 19, 22) 24, 25, 31, 33), 18 pe (822-11, 12 15, 18, 20, 21, 23,

26-28, 30, 32, 34-39), 2 p2 (322-100, 101), 5 1p (322-10, 13, 14,17, 29), 104%,

689, 232 P, 488 L [UCLA]; same data (UCLA 323), 7 lpo’ (323-11, 14, 15, 21,

92,94, 96).42 lp? (323-10, 12,13; 16-20, 23, 25, 27; 29), 1p (823-100), 1 1p

(323-28), 640°, 319, 112 P, 436 L [UCLA].

New York: Buffalo, 5 Sept 1909, M.C. Van Duzee, 1 o [CAS]. Ithaca, 21

Aug-24 Oct 1932, F.C. Baker, 9 ¢, 9 9, 1 P, 20L [CU], 3L [USNM], 1 L

[BMNH]; 3 Apr 1934, FC. Baker, rl E{Cul; 31 Aug-1 Oct 1934, F.C. Baker,

11lpo& (28), 1 lp? (15), 1 p? (20), 3% genitalia, 3 P [CU]; 15 Aug-3 Nov 1935,

F.C. Baker, 2 lpo (17,18), 21p$ (24, 26), 119 (14), 1 1p (22), 30, 299,1P

[Cu], 1 lps (23), 1 1p? (15) [BMNH], 1 1p (20), 1 lp? (21) [USNM].

North Carolina: Charlotte, 8 July-3 Aug 1915, H.P. Barrett, 10, 18

[USNM]. Fayetteville, Fort Bragg, 29 Oct 1943, 3 L [USNM], 1 L [HARM].

Franklin, 8 June 1957, W.R.M. Mason, 1 o [CNC]. Maxton, Nov-Dec 1943,

A.B Klots; 3.26.9 [USNM]; June 1944, K. Snyder, 7°, 59 [UCLA]; 31 Oct

1944, 19 [UTAH]; 6 Aug 1945, 1 L [UCLA], 1 L [UTAH].

Ohio: Canton, 22 Dec 1967, T.J. Zavortink (UCLA 436), 1 L [UCLA]; same

data (UCLA 437), 7 1po' (437-10, 13-17, 19), 3 1p9 (437-11, 12, 18), 14, 32, 7 P,

8 L [UCLA]. Columbus, 17-24 Aug 1942, H.W. Smith, 2 [UCLA]. Toledo, 3

Aug 1915, D.W. Iford, 1 L [USNM].

Oklahoma: Fort Sill, 2 9 [USNM].

Oregon: Corvallis, 27 June 1967, L. F. Lewis, 4 L [UCLA].

Pennsylvania: Roxborough, 15 Aug 1915, G.M. Greene, 1 o [USNM].

South Carolina: Clemson, 20 May 1941, D. Dunavan, 1? [USNM]; 4 June

1945, O.L. Cartwright, 3 L [USNM]. Columbia, 1 Aug 1906, 3? [USNM]; 14

Oct 1944, 1 o [USNM]. Myrtle Beach, 10 July 1943, 2L [USNM]. Sumter Co.,

8 Aug 1944, SoC, Sabrésky, tc [USNM].

Tennessee: Memphis, 2 June 1934, J. A. Le Prince (46), 2 [USNM]. Tip-

tonville, 4 Sept 1928, W.D. Kiser (150), 1 * [USNM]. Walnut Log, Obion Co.,

3 Sept 1935, H. A. Johnson, 1 o [USNM].

Texas: Austin, 10 Mar 1947, O. P. Breland, 1 9 [CNC]; 5 May 1949, O. P.

Breland (10.5), 4 ot, 22 [CNC]; 2 Aug 1957, K. Riemann, 1 L [USNM]. College

Station, Sept 1905, C. E. Sanborn, 1 ¢ [USNM]. Dallas, 17 Apr 1906, F.C. Pratt,

1 [USNM]. Houston, 24 May 1966, R.D. Walsh (TEX 35), 8 L [UCLA]; 23

Feb 1967, R.D. Walsh (TEX 45), 16 L [UCLA]; 5 L [HARM]. Luling, Palmet-

to State Park, 13 Sept 1942, H.R. Roberts and E.S. Ross, 1 L [CAS]; 25 Oct

1942, E.S. Ross, 1L [CAS]. San Antonio, 14-18 June 1942, E.S. Ross, 36,

2 2. 3L [CAS]; 2-3 July 1942, E.S. Ross, 2, 1 9 [CAS]. “Texarkana, Aug

Zavortink: Genus Orthopodomyia 47

1942, E.S. Ross, 1 L [CAS]. No locality, 26 Apr 1943, A.C. Chandler, 2 L [CU].

Utah: Bluff, 13 May 1967, J.H. Arnell and L. T. Nielsen (HA-2-67), 4,

49, 3P, 7L [UTAH]; same data (HA-7-67), 20, 29, 2P, 4L [UTAH]. Can-

yonlands National Park, Salt Creek, 12 May 1967, J.H. Arnell and L. T. Niel-

sen (HA-5-67), 40%, 29,.3 P, 6 L [UTAH].

| Virginia: Bluemont, 27 July 1904, 1 o& [USNM]; 10 Mar 1905, 1 o [USNM].

Falls Church, 17 Sept-8 Oct 1961, W.W. Wirth, 1 0, 1 9 [USNM]. Great Falls,

5 Oct 1906, T. Pergande, 1 [USNM]. Mt. Lake, 16 Aug 1943, O. A. Johann-

sen (2621), 1 L [CU]. Quantico, July 1926, R.L. Turner, 1 2 [USNM]; 1927,

S.S. Cook, 1 o [USNM]. Rosslyn, 2-3 Oct 1903, T. Pergande, 1 o, 1 2? [USNM].

St. Elmo, 4 June, F.C. Pratt, 19 [USNM]. Virginia Beach, 5 May 1928, L.L

Williams, Jr., 12 [USNM]. Williamsburg, Camp Peary, 3 July 1943, G.E.

Bohart, 2, 39 [CAS]; 3 July 1943, R.M. Bohart, 1L [USNM]; Nov 1943, 6 L

[UCD]. Woodstock, 4-16 Aug 1904, F.C. Pratt, 4%, 19, 4 L[USNM].

NO DATA. (233), 39 [USNM]; 6 2 (R-15, 39-1, 72, 125, 149, 188) [USNM];

1L (180-14) [CAS]; 1 o, 19, 1 o genitalia, 2L [USNM], 1 9 [UCLA], 1° [CAS].

Additional records from the literature. CUBA. La Habana: Isla de Pinos,

Nueva Gerona (Montchadsky and Garcia, 1966:36).

Oriente: Baracoa (Montchadsky and Garcia, 1966:36).

HAITI. vide waverleyi.

MEXICO. Tamaulipas: Vargas, 1956: 31).

UNITED STATES. Connecticut: New Canaan, 24 ree 1929, Ri EF. Bart

(Matheson, 1945:27).

Kentucky: Christian, Fayette, Graves, Hardin, McCracken, Meade and Un-

ion Counties (Quinby, Serfling and Neel, 1944:548-549).

Minnesota: Scott and Dakota Counties, 17, 22 Sept 1957, L.R. Abrahamsen,

P, L (Price and Abrahamsen, 1958:92).

North Dakota. vide Carpenter and LaCasse, 1955:101-103.

Rhode Island: Westerly, Great Swamp, 1942 (Knutson, 1943:317).

South Dakota: Bon Homme and Yankton Counties, A (Edman, 1962:431).

VIRGIN ISLANDS. vide Flemings and Walsh, 1966:426.

3. Orthopodomyia alba Baker

Figs. 2,7, 8

1936. Orthopodomyia alba Baker, 1936:1-4. *TYPE: holotype * with associ-

ated larval and pupal skins (7), Ithaca, [Tompkins Co.], New York, United

States, larva found in an elm or maple treehole, 12 Nov 1935 (date of

eclosion), F. Baker [USNM, 51387].

Orthopodomyia alba of Shields and Miles (1937:237); Penn (1938:5); Pennebaker

(1938:4); Shields (1938:429); Edwards (1939:123); King, Bradley and McNeel

(1939:56); Reeves (1941:69, 70, 71,72); King, Bradley and McNeel (1941:61);

Day (1943:30, 31, 32, 37, 39, 40, 42); Gurney (1943:931, 932, 935); King, Roth,

Toffaleti and Middlekauff (1943:574); Bradley, Fritz and Perry (1944:109);

Dorsey (1944:379, 381, 382, 383, 385); King, Bradley and McNeel (1944:61);

Matheson (1944:249); Middlekauff and Carpenter (1944:92); Olson and Keegan

(1944a:781, 782; 1944b:848); Randolph and O'Neill (1944:83); Schoof and Ash-

ton (1944:6-8); Carpenter, Chamberlain and Wanamaker (1945:402); Good

(1945:175); Harden (1945:131); Kitzmiller (1945:409); Peterson and Smith

(1945:381); Carpenter, Middlekauff and Chamberlain (1946:114-116); Jen-

kens and Carpenter (1946:33, 41-43); Miles and Rings (1946:390, 391); Bre-

48 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

land (1947a:185-187; 1947b:81-86); Michener (1947:361); Ross (1947:37);

Weathersbee and Arnold (1947:225); Barnes, Fellton and Wilson (1950:82);

Bohart (1950:403, 404); Darsie (1951:41); Darsie, MacCreary and Stearns

(1951:144); Wilkins and Breland (1951:225-240); Yamaguti and LaCasse

(1951:8-12); Breland (1952:254); Carpenter (1952:252); Yamaguti (1952:16,

17); Hedeen (1953:3,6); Lake (1953:154, 155); Sudia and Gogel (1953:129-

130); Carpenter and LaCasse (1955:98-100); Breeland (1956:101); Pratt

(1956:8); Bickley (1957:24); Breland (1957:305, 307; 1958:219, 221); Burbu-

tis (1958:210, 212); Breland (1959:137-141); Stone, Knight and Starcke (1959:

122); King, Bradley, Smith and McDuffie (1960:135-136); Breeland, Snow and

Pickard (1961:306); Breland (1961:360, 372, 374); Johnson (1961:55-56); Por-

ter, Evans and Hughes (1961:235); Miller (1962:309-310); Tinker and Stojan-

ovich (1962:582); Dodge (1963:804, 808); Warren and Breland (1963:622);

Ross (1964:107); Chapman (1965:435, 438); Ross and Horsfall (1965:14, 24,

43,44); Stone, Sabrosky, Wirth, Foote and Coulson (1965:108); Wiseman

(1965:58); Dodge (1966:378).

FEMALE (fig.2). Wing: 3.35 mm. Proboscis: 2.00 mm. Forefemur:

1.89 mm. Abdomen: about 2.5mm. Head: integument of head capsule, cly-

peus, torus and flagellum brown to black; erect scales dark brown or black;

palpus about 0.35 of proboscis; flagellar segments 1-3, 4 or 5 with scales.

Thorax: integument of mesonotum and scutellum evenly reddish-brown to brown

or black, postnotum usually lighter, light to dark brown, pleuron reddish-

brown to brown or black anteriorly, lighter along most sutures and often pos-

teriorly; supraalar scale line present; lateral prescutellar and inner dorso-

central scale lines not connected; white posterior fossal scales absent; outer

dorsocentral scale line with numerous scales, never curved laterad anteriorly;

lateral lobes of scutellum usually without white scales; lower stp bristles usu-

ally 0-2, weakly to moderately developed, present among uppermost lower stp

scales only; mep bristles all dark; pleural scaling with broad but well defined

lines; ppn scales present; psp scales usually present; stp horizontal scale

lines usually distinct, sometimes joined behind upper stp bristle, stp vertical

scale line not extending dorsad of level of lower edge of mep, some lower and

upper scales usually semierect; lower anterior mep scale patch broad, not

connected to upper. Legs: integument of forecoxa brown to black, midcoxa

lighter, hindcoxa beige to brown; legs with dark-scaling brown or black; fore-

femur with anterior surface streaked in basal 0.05-0.33; midfemur with ante-

rior surface white-scaled in basal 0.05-0.25, posterior surface light-scaled

in basal 0.05-0.5; hindfemur with anterior surface light-scaled in basal 0. 05-

0.3; hindtibial anterior, dorsal and posterior surfaces speckled with white

scales; foretarsus entirely dark-scaled; midtarsal segment 1 with complete or

ventrally incomplete basal white ring and usually with dorsal apical white patch,

segment 2 sometimes with small dorsal white patch at base and rarely with one

at apex; hindtarsus with apex of segment 1 and base and apex of segments 2 and

sometimes 3 with complete white ring, segment 5 all white dorsally. Wing: light

spot in area of crossveins usually well developed and conspicuous to unaided eye,

white patches on veins M and Cu, usually large, but vein R4,5 usually dark-

scaled at base and white patch on vein R943 reduced or absent. Haltere: integu-—

ment of stem whitish to beige, knob beige to brown. Abdomen: integument of

tergite I whitish to brown, sternites II-VII whitish to brown, distal segments

darker, sternite VIII tan to brown; abdominal segments with dark scales brown

or black; tergite II largely white-scaled, dark scales usually present in apico-

lateral patch, sometimes in narrow apical band; tergites III-VIII with basolater-

Zavortink: Genus Orthopodomyia : 49

al white patch and sometimes with irregular, narrow, beige basal band; sternite

II entirely white-scaled or with some apical scales dark; sternites II-VII white

basally, dark scales in apicolateral patch or in apical band, distal segments with

dark-scaling more extensive; sternite VIII with dark scales laterally.

MALE. Essentially as infemale. Labium: as in female or with white

scale lines broadened just distad of level of distal end of palpal segment 2.

Palpus: about 1.01 of proboscis. Wing: white scales on veins behind vein R1

sometimes dingy and difficult to distinguish from dark scales. Abdomen: as in

female or with tergites II-VII with rather distinct white basal band; sternites

II- VII usually darker than in female, sometimes all dark-scaled; tergite VIII

scales all dark or mixed dark and white, sternite VIII with white apical and

basal bands. ; :

MALE GENITALIA (fig. 7). Segment VIII: tergite lobe longer than broad,

narrowing distally, apex emarginate or very irregular. Segment IX: tergite

with 2 or 3 bristles on each side. Sidepiece: basal mesal lobe with 3 or 4 stout

bristles which are straight or only very slightly bent mesally near apex and

with 2 or 3 finer bristles. Aedeagus: ventral spines variable in position; pre-

apical ridge with 2-5 processes. Proctiger: paraproct sclerotization with 2-4

apical teeth; cercal setae 2-4.

PUPA (fig. 7). Abdomen: 3.5mm. Trumpet: 0.5mm. Paddle: 0.8 mm.

Cephalothorax: hairs 8,9-C never as long as trumpet. Trumpet: deep tan to

brown except at extreme base and apex and largely darker than darkest part of

cephalothoracic integument. Metanotum: hair 12-C usually moderately devel-

oped. .Abdomen: hairs 6, 7-I short; 1-II moderately developed, usually 3b (1-

6); 2-II-V usually 0. 3-0.4 length of hair 1 of corresponding segment, 2-VI at

least 0.5 length of 1-VI; 6-II short and fine, similar to 7-II; 5-III usually dou-

ble (single, double), developed similarly to 5-IV-VI, and usually more strongly

developed and longer than 3-III; 10-III-VII not extending more than 0.5 way to

apex of following segment; 5-IV, V extending to about 0.5 of following segment.

Paddle: usually more or less oboval in outline; extreme apex truncate or emar-

ginate; inner part without minute spicules near inner margin.

LARVA (fig.8). Head: 0.82 mm. Siphon: 0.63 mm. Anal Saddle: 0.38

mm. Most branched hairs with conspicuous barbs. Head: largely light straw

colored, cephalic edges darker, collar brown; mental plate tan, usually with

7-9 (7-11) teeth on each side; hair 2-C absent; 9-C usually 3-5b (2-7), branches

strong; 11-C moderately developed, usually 7, 8b (4-16); 13-C strongly devel-

oped, usually 7, 8b (5-12). Antenna: largely tan colored, apex lighter. Thorax:

epidermal pigment absent; spicules not conspicuous; hairs strongly pigmented;

hair 0-P usually 3, 4b (2-5); 6-P sometimes multiple; 1-M strongly developed,

usually double (1-3b); 1-T strongly developed, usually single (1-3b); 13-T

strongly developed, usually 5-8b (2-8). Abdomen: segments VI and VII never

with sclerotized plates; hairs strongly pigmented; hair 1-I strongly developed,

long, usually single or double (1-3b); 4-I longer than 3-I; 6-I usually 3b (2-5);

7-I usually 3, 4b (3-6); 1-II strongly developed, long, usually single or double |

(1-3b); 4-II well developed; 5-II-VI well developed, usually 3b (2-4); 6-II usu-

ally 2, 3b; 7-II strongly developed; 1-III long, usually single (single, double);

13-III-V usually 2, 3b (2-6), and usually longer than hair 10 of corresponding

segment; 1-VI, VII moderately developed; 13-VII usually 3, 4b (2-4). Segment

VII: sclerotized plate never present; anterior comb scales usually 12-14 (10-

17), posterior comb scales usually 8-11 (5-11); posterior row of comb scales

about 0. 84-1.07 length of anterior row; hair 3-VIII weakly to moderately devel-

oped, usually 2, 3b (1-5). Siphon: index about 2.6-3.4; base light brown in

color, darker brown or blackish towards apex; integumentary sculpturing ab-

50 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

sent or very weak, imbricate; hair 1-S located near base of siphon, usually 3,

4b (2-6), weakly to moderately developed and with longest branch usually not

extending much beyond middle of siphon. Anal Segment: saddle complete or

incomplete; saddle largely tan, dorsal and apical portion darker, brown; in-

tegumentary sculpturing inconspicuous, imbricate; diameter of saddle or anal

segment at most proximal point considerably smaller than diameter just basad

of ventral brush, so that dorsal and ventral surfaces diverge distally when

viewed from the side and diameter of saddle or anal segment at most proximal

point usually equal to or greater than diameter of base of siphon; hair 1-X usu-

ally 2, 3b (1-3), branches coarse; ventral brush usually with 6 pairs of hairs;

more strongly developed hairs of ventral brush usually with 4-8 branches; gills

very short, rounded apically, dorsal pair only slightly longer than ventral pair

and 0.5-1.0 length of anal saddle. :

BIONOMICS. Larvae have been collected in treeholes and artificial con-

tainers. They have usually, but not always, been found in association with sig-

nifera. Breland (1947:185) found live larvae and pupae in the moist debris of a

treehole with no free water.

-Adults have been caught in light traps and found resting in treeholes and

barns. Sudia and Gogel (1953:130) state that a female fed at night on a chick.

Additional information on the bionomics of alba is given in the Biology and

Ecology section.

SYSTEMATICS. Alba is one of the 2 sympatric species known in the Signi-

fera group. It occurs with signifera throughout the eastern United States, in

northeastern Mexico and as far west as New Mexico.

Adults of alba and signifera are very similar. While some authors (see

Baker, 1936:7; Carpenter and LaCasse, 1955:98-99) have provided a few char-

acters for separating the adults of these species, the characters have not been

found reliable enough to permit separation of individual specimens and as a

consequence most workers have considered the adults to be indistinguishable.

Probably the best character was Baker's observation that the cuticle of the first

abdominal tergite of signifera was purplish-brown whereas that of alba was

yellowish. The purple epidermal pigment of the signifera larva is also present

in the pupa and adult. While it is normally obscured by the dark integument in

these stages, it is quite obvious in teneral individuals and is sometimes visible

through the largely unscaled first abdominal tergite of pinned specimens of sig-

nifera. Two very reliable characters for separating the adults of alba and sig-

nifera have been found in this study. The better one is the number of lower stp

bristles. In alba there are seldom more than 2 bristles on stp below the level

of the lower edge of mep and these bristles are usually present only among the

uppermost lower stp scales. In signifera 4 to 10 bristles are present and they

are found throughout the length of the lower stp scales. The white spot over

the crossveins of the wing is usually conspicuous in both alba and eastern sig-

nifera, but in alba the base of vein R445 is normally dark-scaled whereas in

signifera it usually has a patch of white scales.

Alba is the only species in the Signifera group which has very distinctive

immature stages. The pupa differs from that of all other species in the devel-

opment of hairs 5-III and 2-II-VI. There are numerous additional characters

(see Darsie, 1951:40-41) for separating the non-hairy pupa of alba from the

hairy pupal form of signifera which occurs in all areas where the 2 species are

sympatric. The fourth instar larva of alba is very different from that of any

other species. The differences are largely the result of 2 phenomena; first,

the larva is neotenic, that is, it retains many features in the fourth instar that

are found in the third instar larvae of other species, and second, numerous

Zavortink: Genus Orthopodomyia ol

elements of the chaetotaxy are genetically fixed at a high level of hairiness.

Neotenic features which distinguish alba are the development of hairs 6-I, II,

3-VIII, 1-S and 4-X and the complete lack of abdominal plates on segments VI-

Vill. Particularly diagnostic setae which are fixed at a high level of hairiness

are 11 and 13-C. Other diagnostic features of the alba larva, such as its com-

plete lack of epidermal pigment and the anterior row of comb scales being

Short, are not the result of either neoteny or hairiness.

Except for the reduced pale-scaling in depauperate specimens, especially

males, the species appears to be quite uniform in all stages throughout its

range.

Largely on the basis of the few lower stp bristles, I am considering alba to

be more primitive than either signifera or waverleyi. In general, evolution in

the Signifera group seems to be in the direction of increasing numbers of tho-

racic bristles and extent of white thoracic and wing scaling. In particular,

while the more primitive Kummi and Pulchripalpis subgroups have no lower stp

bristles at all, signifera and waverleyi have a large number of them. Alba

would appear, then, to be a species representing a stage in the evolution of the

more advanced signifera and waverleyi. However, because of its apomorph

larva, alba cannot possibly be ancestral to the signifera-waverleyi stock; the

unique alba larva clearly had to have arisen after the isolation of alba from the

line culminating in signifera and waverleyi. I believe that the original separa-

_ tion of the alba and signifera-waverleyi stocks was geographical, possibly in

different glacial refuges, and that not only isolating mechanisms but also at

least a portion of the alba larval differences evolved before the subsequent re-:

contact of the lines. While it is true that the larva of alba is very different

from those of the other species, since most of its morphological differences

are the result of neoteny and genetically fixed hairiness, it could have arisen

extremely rapidly. If there had been no larval differentiation between the stocks

when they initially reestablished contact, one would have probably eliminated

the other by competition. But, as natural selection would tend to preserve any

additional differences reducing competition between sympatric species, some

of the features of the alba larva could have arisen after sympatry with the sig-

nifera-waverleyi line. At any rate, I feel that the manifold differences of the

alba larva, which must enable it to occupy a different niche in the treehole

habitat, are what permit the coexistence of alba and signifera over a broad

geographical area today.

DISTRIBUTION. New Mexico, Missouri and New York, south to Coahuila,

Texas and Georgia. Material Examined: 732 specimens; 86 0, 789, 179 P,

389 L; 42 individual rearings (39 larval, 3 incomplete).

MEXICO. Coahuila: Allende, 22 Aug 1961, J. Delgado, 2 L [ISET].

UNITED STATES. Alabama: Wilson Dam, 3 July 1936, S.E. Shields (4-

248), 1 [USNM]; 16 July 1936, S.E. Shields (1-261), 19, 1 L [USNM].

Arkansas: Fort Smith, Camp Chaffee, 21 Sept 1943, B. Crafchick, 1 ¢

[CU]. Stuttgart, 28-29 Apr 1915, J.A. Le Prince, 1 9 [USNMI].

District of Columbia: 20 July 1901, N. Banks, 1 o [USNM]; 4 Sept 1942,

1 o [USNM]; 26 June 1943, 19 [USNM]; 9 July 1943, 1 [USNM]; 20 Aug-25

Sept 1943, Io’, 22 [USNM]; C.N. Ainslie, 12 [USNM]; W.V. Warner, 1% |

[USNM].

Georgia: Atlanta, 25 Mar 1953, W.D. Sudia and R. H. Gogel, 30°, 39, 6L

[UCLA], 4 L [USNM].

Maryland: College Park, 1 2 [USNM].

Missouri: Neosho, Camp Crowder, 6 Oct 1942, A.B. Gurney (225), 30,

——————

6L [USNM]; 9 Oct 1942, A. B. Gurney (239), 10° [USNM]; Oct 1942, A. B. Gur-

o2 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

ney, 1 P, 3 L[USNM], 1 P, 3 L [UCD].

New Jersey: Newport, 19 July 1937, J. A. Rowe, 1 2 [USNM].

New Mexico: Alcalde, 6 L [HARM].

New York: Ithaca, 7-16 Sept 1934, F.C. Baker, 21lpo% (1,2), 112 (4), 2P

6 L [CU], 1 L [BMNH], 1 L [USNM]; 5 Feb 1935, F.C. Baker, 1 P [CU]; 17

Aug-12 Nov 1935, F.C. Baker, lpo holotype (7), 2 lpo (1,2), 2 lp? (8, 9)

[USNM], 1 1p? (5), 39, 1P [CU], 1 lpo (4), 1 lp? (6) [BMNH]; 13-16 Aug 1936,

F.C. Baker, 119 (18), 1 o [CU]. |

Ohio: Canton, 22 Dec 1967, T.J. Zavortink (UCLA 436), 14 lpo (436-20-

22, 24-30, 33, 34, 36, oy), O1pe (436-23, 31, 32, 35, 38), 1 1p (436-37), 39 P, 241 L

[UCLA]; same data (UCLA 437), 8 1lpo (437-20-26, 28), 2 1p? (437-27, 29), 1,

5 P, 60 L [UCLA]; same data, mixed collections (UCLA 436 and UCLA 437),

Slo, 372, 89:-P (UCLA).

Oklahoma: Oklahoma City, 11 July 1934, 192 [USNM].

Texas: Austin, 16 June 1947, O.P. Breland. oo, 69 |USNM|, 4:0, 29

[CNC]; 8 July 1947, O. P. Breland, 1, 29 [CNC]; 30 Nov 1949, O. P. Wilkins,

3 L [UCD]; 26 Aug 1957, K. Riemann, 1 L [USNM]. Brownsville, 20 Aug-24

Nov 1925; R.L. Turner, 3 0, 1°9 [USNM]. Kerrville, 1 Aug-4 Sept 1953, L.J.

Bottimer, 1 o, 19 [USNM]. San Antonio, 1L [HARM]. Sonora, May 1934, 1 ?

[Cul

Virginia: Williamsburg, Camp Peary, Nov 1943, 2 L [UCD]. :

Additional larval records from the literature. UNITED STATES. Illinois:

Onarga, L (Ross, 1947:37).

Kentucky: Fort Knox, 22, 24 Aug 1944, J.B. Kitzmiller and T.E. Potts,

L (Kitzmiller, 1945:409).

Louisiana: New Orleans, 23 Nov 1937-May 1938, L (Pennebaker, 1938:4).

Mississippi: Hattiesburg, Camp Shelby, 10 Aug 1943, L (Michener, 1947:

361).

North Carolina: Manchester, 18 Sept 1942, S.C. Schell, L isehoot and Ash-

ton, 1944:6-8).

Tennessee: Knoxville, 11 July 1955, S.G. Breeland, L (Breeland, 1956:

101).

PULCHRIPALPIS SUBGROUP

4. Orthopodomyia pulchripalpis (Rondani)

Figs. 2, 9, 10

1872. Culex pulchripalpis Rondani, 1872:31. *TYPE: lectotype © with genitalia

slide 670327-1, Italy; PRESENT SELECTION [FM].

1919. Orthopodomyia albionensis MacGregor, 1919:451-454. *TYPE: lectotype

oO, Epping Forest, Essex, England, reared from a larva found in a beech

treehole, July 1919 (date of eclosion), A. Macdonald; PRESENT SELEC-

TION [BMNH]. Synonymy with pulchripalpis by Edwards (1921:290, 291).

Orthopodomyia pulchripalpis of Seguy (1920b:253); Edwards (1921:290-291);

Seeuy (1921a:182, 187; 1921b:110, 112-114); Dyar (1922:67); Martini (1922:

99); Dyar (1923b:96); Seguy (1923:64, 98, 130, 154-155); Keilin (1927a:370;

192'7b:196); Stackelberg (1927:68-69); Beattie and Howland (1929:49, 50, 56);

Bashkareva (1931:56, 58); Brighenti (1931:219); Kalandadze (1931:113);

Velichkevich (1931: 316, 320-321); Edwards (1932:107, 108); Keilin (1932:280,

281); Macan and Tutin (1932: 283); Tate (1932:111- 120): Marshall and Staley

Zavortink: Genus Orthopodomyia D3

1933:435); Edwards and James (1934:20); Marshall and Staley (1935:531);

Pandazis (1935a:926; 1935b:3, 18); Montchadsky (1936:218-222); Stackelberg

(1937:114-115); Callot (1938:152); Marshall (1938:258-264); Stephanides

(1938:251); Edwards (1939:122); Edwards, Oldroyd and Smart (1939:1, 22-

23); Macan (1939:266-267); Clastrier (1941:443-446); Cambournac (1943:71-

77); Callot and Ty (1944:65); Cambournac (1944:256); Cafiamares (1945:838,

842, 849); Classey (1946:113); Shillito (1948:25, 26, 27); Bohart (1950:403):

Mattingly (1950:113, 117); Montchadsky (1951:143-146); Aitken (1954:465);

Peus (1954:79-80); Beam ci) Andarelli and Duzer (1954:274); Clastrier (1955:

274-275); Doby (1955:370); Doby and Doby-Dubois (1955:384); Mattingly

(1955:32); Mihalyi (1955:14); Hedeen (1958:320); Mihalyi (1959:142-143);

Parrish (1959:266); Rioux and Nicoli (1959:74); Senevet and Andarelli (1959:

301-309); Stone, Knight and Starcke (1959:123-124); Coluzzi (1962:14); Rioux,

Juminer and Kchouk (1964:14, 18); Callot and Rioux (1965:243); Dubose and

Curtin (1965:349, 352); Bailly-Choumara (1966:39-41); Bozhkov (1966:571,

572,573); Dodge (1966:379).

Orthopodomyia (Bancroftia) pulchripalpis s of Shannon and Hadjinicolaou (1937:

173,174, 175, 176, 178); Rioux and Arnold (1955:278); Rioux (1958:257-260).

Orthopodomyia albionensis of Lang (1920:109-111); Seguy (1924:201; 1925:78);

Martini (1930:194-196).

Culex pulchripalpis of Ficalbi (1896:256; 1899:220-222); Giles (1900:275-276;

1902:426); Husson (1908:86, may refer to Aedes).

FEMALE (fig. 2). Wing: 4.95 mm. Proboscis: 2.82 mm. Forefemur:

2.97 mm. Abdomen: about 3.5mm. Head: integument of head capsule, cly-

peus, torus and flagellum dark brown or black; vertex and occiput with white

decumbent scales usually more numerous laterally: erect scales dark brown

or black; dorsolateral broad scales dark and white; clypeus an elongate, some-

what angular lobe, bare; labium largely dark-scaled, with white ring at about

0.67 distance from base; palpus about 0.44 of proboscis, 5-segmented, seg-

ment 5 minute, segment 4 short, segments 1, 2 and 3 largely dark-scaled, usu-

ally with dorsal white spot at base of segments 2 and 3 and sometimes at apex

of segments 1 and 2; flagellar segment 1 with brown scales. Thorax: integu-

ment of mesonotum, scutellum, and postnotum evenly reddish-brown to black,

pleuron reddish-brown to black anteriorly, usually lighter behind and below

ppn, in posterior portion of stp and mep, and on metapleuron and metameron;

paratergite narrow; supraalar scale line present; lateral prescutellar and in-

ner dorsocentral scale lines not joined; posterior fossal white scales absent;

outer dorsocentral scale line extending to scutellum; lateral lobes of scutellum

without scales; lower stp bristles absent, so that there are no bristles among

lower stp scales; upper posterior mep bristles dark in color, anterior mep

bristles 0-3, weakly developed, dark in color; pleural scaling with lines rather

broad; ppn n scales present; psp scales absent; pst with scales present only near

ppl; stp with horizontal scale lines usually not connected, although lower is

quite broad; upper stp scale line sometimes more or less connected to scales

on pra; vertical stp scale line not extending dorsad of level of lower edge of

mep and without semierect scales; lower anterior mep scale patch broad,

sometimes contiguous with upper. Legs: integument of forecoxa reddish-brown

to dark brown, midcoxa somewhat lighter, hindcoxa much lighter; all coxal

scales white or fore- and midcoxae with lower scales brown; trochanters with

all scales white or with some brownish; remaining leg segments with dark

_ scales brown or black; forefemur with anterior surface usually streaked with

white or dingy-white scales in basal 0.10 to 0.67 dorsally, remainder dark-

o4 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

scaled, ventral margin speckled with white scales from base to apex, posterior

surface usually largely white-scaled, dark scales in longitudinal streak from

base to near apex along or near ventral margin, usually along dorsal margin |

apically, and at apex; midfemur with anterior surface largely dark-scaled with

white speckles, sometimes white speckles concentrated inmoreor less distinct

median or ventral longitudinal streak basally, posterior surface usually pale-

or white-scaled in at least basal 0.33 and to or near apex in a progressively

narrowing streak, remainder dark-scaled with white speckles; hindfemur with

anterior surface usually pale- or white-scaled at least dorsally in basal 0.10

to 0.67, remainder entirely dark-scaled or with white speckles which may be

concentrated near dorsal margin, posterior surface usually pale- or white-

scaled in basal 0.10 to 0.67, remainder entirely dark-scaled or with white

speckles; tibiae largely dark-scaled; foretibial anterior surface with white

scales in line or line of speckles from base to or near apex, posterior surface

with irregular speckling or mottling of white scales basally; midtibial anterior

and posterior surfaces usually and dorsal surface sometimes with irregular

speckling or mottling of white scales basally or from base to near apex;. hind-

tibial anterior and dorsal surfaces with speckled white scales in at least cen-

tral portion of segment; tarsi largely dark-scaled; foretarsal segment 1 some-

times with small dorsal white patch at base and less often with smaller one at

apex; midtarsal segment 1 with small dorsal white patch at base and sometimes

with one at apex, segment 2 less often with very small dorsal white patch at

base and rarely with one at apex; hindtarsal segment 1 with posteriorly incom-

plete basal white ring, segments 1 and 2 sometimes with complete apical ring,

segment 5 entirely white-scaled dorsally. Wing: largely brown- or black-

scaled, white scales in short line about twice length of remigium at base of

vein R, usually speckled on vein Sc, and sometimes in very small patch at base

of costa; veins behind vein Ry with very narrow scales; scales on vein 1A lying

parallel to vein. Haltere: integument of stem whitish to beige, knob whitish to

tan, scales medium wide. Abdomen: integument of tergite I and sternites I-

VII whitish to brown, apex of sternites often darker than base and more distal

sternites often darker than proximal ones, integument of sternite VIII tan to

brown; abdominal segments with dark scales brown to black; tergites TI-VII

with conspicuous broad, basal, white bands, apex dark-scaled; tergite VIII

_ dark-scaled with white scales basally or basolaterally; sternite II entirely

white-scaled; sternites HI-VII white-scaled basally, dark-scaled apically or

apicolaterally, dark-scaling becoming more i cgecaonel on distal segments;

sternite VIII bare.

MALE. Essentially as in female. Labium: largely dark-scaled, usually

with small ventral white spot or complete narrow white ring just distad of level

of distal end of palpal segment 2. Palpus: about 0.99 of proboscis, segment 3

with well developed apical bristles; largely dark brown- or black-scaled, seg-

ments I and 2 white at apex dorsally, segments 2 and 3 white at base dorsally.

Antenna: flagellar segment 1 with black or brown scales dorsomesally. Tho- |

rax: upper posterior mep bristles not reduced in number. Abdomen: segment

VIII entirely white-scaled or with a few dark scales.

MALE GENITALIA (fig. 9). Segment VIII: tergite lobe large, sides more

or less parallel or slightly diverging distally, apex usually truncate, with ir-

regular edge, sometimes with small or large emargination. Segment IX: ter-

gite with 2-4 bristles on each side, sternite often with small projection in mid-

dle of cephalic edge. Sidepiece: basal mesal lobe with 4-6 usually straight,

stout, bristles and 1-3 finer bristles. Clasper: not strongly constricted distad

of origin of apodeme, but tapering gradually from base to apex. Aedeagus:

Zavortink: Genus Orthopodomyia D0

more or less pyriform or sometimes with lateral margins from or near apex

to broadest point of aedeagus nearly straight; ventral triangular spines usually

distad of middle of aedeagus; preapical ridge with 1-4 processes. Proctiger:

paraproct sclerotization with 3 or 4 apical teeth; cercal setae usually 3-6.

PUPA (fig.9). Abdomen: 4.5mm. Trumpet: 0.7mm. Paddle: 0.9 mm.

Cephalothorax: hairs 8, 9-C never as long as trumpet. Trumpet: uniformly

pigmented or with extreme base somewhat lighter, largely bright amber to tan

and normally concolorous with or lighter than darkest portion of cephalothorax.

Metanotum: hair 12-C usually moderately developed. Abdomen: 6, 7-I usually

medium-long; 1-II moderately developed, usually double (2-7b); 2-II-VI usual-

ly less than 0.3 length of hair 1 of corresponding segment; 6-II short and fine,

Similar to 7-II; 5-III usually double and not as strongly developed as 5-IV-VI

or 3-III; 10-III, IV not extending beyond 0.4 of following segment, 10-V-VII not

extending beyond 0.8 of following segment; 5-IV, V extending to about 0.75 of

following segment. Paddle: usually more or less oval in outline; apex rounded

to slightly truncate; inner part without spicules near inner margin.

LARVA (fig. 10). Head: 1.04 mm. Siphon: 0.97 mm. Anal Saddle: 0.46

mm. Not all branched hairs with conspicuous barbs. Head: largely light brown

colored, lighter cephalad and around imaginal eye, collar darker; mental plate

tan, usually with 8-10 teeth on each side; hair 2-C sometimes present; 9-C

usually single or double (1-3b), fine; 11-C weakly developed, usually with 4-

6b; 13-C moderately developed, usually 3b (2-4). Antenna: uniformly light

straw-colored. Thorax: epidermal pigment weakly developed, pink, violet or

reddish in color; spicules not conspicuous; hairs moderately pigmented; hair

0-P usually with 3-6b (3-8); 6-P single; 1-M moderately to strongly developed,

single; 1-T weakly to moderately developed, usually single or double (1-3b);

13-T moderately developed, 3,4b. Abdomen: segment VII usually with sclero-

tized plate and segment VI often with plate; hairs moderately pigmented; hair

1-I weakly developed, usually 2-4b (1-5); 4-I shorter than 3-I; 6-I usually 5, 6b;

7-I single; 1-II weakly developed, usually double (1-3b); 4-II moderately devel-

oped; 5-II-VI weakly to moderately developed, usually 2, 3b (1-3); 6-II usually

5, 6b; 7-II moderately developed; 1-III weakly to moderately developed, single;

13-III-V usually 2, 3b and shorter than hair 10 of corresponding segment; 1-VI

weakly to moderately developed, usually 2, 3b (1-3); 1-VII moderately devel-

oped; 13-VII usually 5, 6b (5-8). Segment VIII: large sclerotized plate pres-

ent; anterior comb scales usually 16-21 (15-23), posterior comb scales usual-

ly 6-8 (5-9); posterior row of comb scales about 0. 47-0. 67 length of anterior

row; hair 3-VIII strongly developed, usually 5-8b. Siphon: index about 3. 6-

5.0, usually greater than 4.0; largely light brown to brown in color, extreme

base darker, extreme apex lighter; integumentary sculpturing absent; hair 1-S

located 0.4-0.5 from base of siphon, usually 9-13b, strongly developed and

_with longest branches extending to apex of siphon. Anal Segment: saddle com-

plete, light brown; integumentary sculpturing very weakly developed, imbri-

cate; diameter of saddle at most proximal point not or only slightly smaller

than diameter at the most distal point along ventral surface, so that dorsal and

ventral surfaces are parallel or diverge only slightly when viewed from the

side and diameter of saddle at most proximal point usually equal to or greater

than diameter of base of siphon; hair 1-X single, fine; ventral brush usually

with 7 pairs of hairs; more strongly developed hairs in ventral brush with 9-11

branches; gills moderately long, dorsal pair about twice as long as ventral

pair and 0.9-1.3 length of anal saddle.

BIONOMICS. Larvae have been found only in treeholes. Adults have ap-

parently never been collected.

06 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

Tate (1932:111-120) found that reared females would not bite men, birds,

toads, earthworms or slugs and Cambournac (1943:71-77) reported that they

would not take blood from men, birds, rabbits or frogs.

SYSTEMATICS. This is the only Palearctic representative of the Signifera

group. The adult is distinctive enough for the species to be placed in a mono-

typic subgroup. It differs most conspicuously in having the outer dorsocentral

scale line extending to the scutellum, the scales on vein 1A very narrow, vein

R white-scaled at base for a distance about twice the length of the remigium,

the proboscis of the female white-ringed and in lacking white scales on the

basal flagellar segments.

Only non-hairy forms of both the larva and pupa are known for this species.

The larva is difficult to distinguish from signifera from the eastern United

States and the Greater Antilles, and from kummi and waverleyi. In pulchri-

palpis the siphon is usually slightly longer, the dorsal and ventral surfaces of

the anal segment are more frequently parallel, hair 13-VII is usually more

strongly developed and the sclerotized parts are usually lighter colored than in

these other species. The pupa is most similar to that of signifera from Calif-

ornia and the Greater Antilles and waverleyi; it is usually told from these by

its lighter trumpet color and in addition is separated from Californian signifera

by the paddle being narrower. It differs from signifera from the eastern United

States and from kummi in being less hairy.

The femoral ornamentation is quite variable in this species. Since all in-

dividuals in one collection usually have the same pattern and this pattern may

differ from that of individuals from another collection in the same area, this

variation must be environmentally induced. Clastrier (1941:443-445) indicated

that pulchripalpis from Algeria lacked the white proboscis ring in the female

and Bailly-Choumara (1966:39-41) reported the same for material from Moroc-

co. The immature stages from England are slightly less hairy than others

which I have seen.

This species bears a greater resemblance to kummi than to any other spe-

cies in the Signifera group. It is apparently one of the older species in the

group. Some of its peculiarities have evidently been obtained by introgression

from the Oriental Albipes group. In particular, the ringed proboscis and 5-

segmented palpus of the female and the strongly developed lateral prescutellar

scale line are features of pulchripalpis which are found in the Albipes group

but not in other species in the Signifera group. While pulchripalpis is not known

to contact any species of the Albipes group at present, it may have been sym-

patric with some species of that group in southwestern Asia during more mesic

times.

DISTRIBUTION. England, France, Hungary and Ukrainian, south to Mo-

rocco, Tunisia, Turkey and Georgian. Material Examined: 472 specimens;

900, 862, 141 P, 155 L; 40 individual rearings (30 larval, 10 incomplete).

ALBANIA. Mamuras, 13 June 1932, D.J. Lewis, 40°, 42 [BMNH].

ENGLAND. Essex: Epping Forest, 20 July 1920, F.W. Edwards, 1 &

[BMNH]; July 1919, A. Macdonald, & lectotype of albionensis, 1 9 [BMNH];

Aug 1919, F.W. Edwards, 1? [BMNH]; Aug 1919, H. Main, 3 L [BMNH]; Sept

1919, H. Main 1 L [BMNH]; 12 Sept 1919, 1 L [BMNH]; 4 P, 3 L [BMNH].

Loughton, Epping Forest, 12 June 1966, T.J. Zavortink (UCLA 312), 9 lpc&

(312-24-29, 32, 34, 38), 9 lp? (3812-10-13, 15, 16, 20, 35, 36), 6 1p (312-14, 23, 30,

31, 33, 37), 31%, 309, 83 P, 56 L [UCLA]. Wake Arms, Epping Forest, 18

Aug 1919, F.W. Edwards, 1 L [BMNH]. | |

Hampshire: Hayling Island, 1c, 2P, 5L [BMNH]. New Forest, Buskett's

Wood, June 1939, F.W. Edwards, 110, 92, 11 L [BMNH], 10, 1 9 [UCLA].

Zavortink: Genus Orthopodomyia sy

London: Buckingham Palace Gardens, 30 Mar 1926, F.W. Edwards, 1 P,

1L [BMNH]; June 1926, F.W. Edwards, 20°, 3° [BMNH]. Kensington Gardens,

30 Mar 1926, F.W. Edwards, 5 P, 6 L [BMNH]; 10 June 1966, T.J. Zavortink

(UCLA 311), 41po (311-10, 13,17, 21), 81p? (311-12, 14, 16, 18, 23-26), 41p

(311L-14,.15; 20,.22),. list, 4B, 10: IUCLAL.

No locality or date: J. F. Marshall, 30, 29 [USNM], 1, 1 2? [CNC].

GREECE. Macedonia: Lahanas Hills, July-Oct 1935, R.C. Shannon and

J. Hadjinicolaou, 10°, 6L [USNM]. Struma Valley, June-Oct 1934, R.C. Shan-

non, 110, 119, 3 L [USNM]; June 1934, R.C. Shannon and J. Hadjinicolaou,

1 o« [USNM]; Aug 1935, R.C. Shannon and J. Hadjinicolaou, 1, 4? [USNM].

ITALY. Latium: Esperia, 28 May 1960, 30, 12, 2L [UCLA]. Ponte-

corvo, 28 May 1960, 1 P[UCLA]. Castnocielo, 29 Mar 1961, 1 P [UCLA].

Sardinia: Gadoni, 11 Mar 1948, 2 L [BMNH].

No No locality or: date: C. Rondon (3022), o& lectotype of pulchripalpis, 1

[FM].

PORTUGAL. Estremadura: Aguas de Moura, 1937, F.J. Cambournac, 1c,

1° [BMNH]. —

YUGOSLAVIA. Croatia: Zagreb, Pavicic, 2 L [BMNH].

NO LOCALITY. June 1928, Medjid (5214), 2 L [BMNH] (possibly from

Turkey, vide Mattingly, 1955: 32),

Additional records from the literature. ALGERIA. Mouzaia Mountains,

L, A (Clastrier, 1941:443-446).

BULGARIA. Black Sea coast (Bozhkov, 1966:571, 572, 573).

ENGLAND. Buckinghamshire: Burnham Beeches, 1928-1929, M.V.F. Beat-

tie and L.J. Howland (Beattie and Howland, 1929:49, 50, 56).

Cambridgeshire and the Isle of Ely: Cambridge, L (Keilin, 1927a:370).

Surrey: Ripley, 1930, M.E. MacGregor (Marshall, 1938:258-264).

FRANCE. Aude: Sigean, L (Rioux, 1958:257-260).

Bouches-du-Rhone: Saintes-Maries-de-la-Mer, Mar 1955, J.A. Rioux and

J. Callot, L (Rioux, 1958:257-260).

Charente Maritime: La Rochelle, Mar 1955, R.A. Hedeen, L (Hedeen, 1958:

320).

Corsica: Morosaglia and Porto Vecchio, 1, 5 Nov 1959, L (Rioux and Nico-

li, 1959:74).

Gard: Alex, L (Rioux, 1958:257-260).

Herault: Montpellier (Rioux and Arnold, 1955:278).

Indre-et-Lorie: Richelieu, Mar, July 1937, L (Callot and Ty, 1944:65).

Pyrenees-Orientales: Massane, 20 Aug 1953, L (Doby, 1955:370).

Seine-et-Oise: St. Germain, Forét de Marly, M.P. Lesne, L (Seguy, 1920b:

2oa)s

GREECE. Ionian Islands: Corfu (Stephanides, 1938:251).

Thessalia: Tempi Valley, L (Pandazis, 1935a:926).

HUNGARY. Veszprem: Lake Balaton, Badeortes Heviz, 1938,1939, F.

Mihalyi, L (Mihalyi, 1959:142-143).

MOROCCO. Mamora, Mar 1965, L (Bailly-Choumara, 1966:39-41).

SPAIN. Cuenca: Alberca de Zancara, Apr, May 1945, L (Cafiamares, 1945:

838, 842, 849).

erane: Gerona, L (Rioux, 1958:257-260).

TUNISIA. Kroumirie Mountains, 27, 28 Mar 1964, L (Rioux, Juminer and

Kchouk, 1964:14, 18).

TURKEY. (Parrish, 1959: 266). ;

UNION OF SOVIET SOCIALIST REPUBLICS. Adygey: Sochi, Bashkareva

(Stackelberg, 1937:114-115).

58 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

Adzhar: Lindrop (Stackelberg, 1937:114-115).

Crimean Oblast: (Velichkevich, 1931:316, 320-321).

Georgian: Abkhaz (Kalandadze, 1931:113).

KUMMI SUBGROUP

FEMALES. Head: vertex and occiput with few white decumbent scales;

dorsolateral broad scales dark and white; clypeus more or less triangular,

bare; palpus 0.41 to 0.45 of proboscis, 4-segmented, segment 4 very short,

entirely white-scaled; flagellar segments 1-4, 5 or 6 with white scales. Tho-

rax: paratergite moderately broad; supraalar scale line present; lateral pre-

scutellar and inner dorsocentral scale lines not joined; posterior fossal white

scales absent; outer dorsocentral scale line not reaching scutellum; lateral

lobes of scutellum without scales; lower stp bristles absent, so that there are

no bristles among lower stp scales; upper mep bristles dark in color, anterior

mep bristles absent; pleural scaling with all lines or patches very narrow; ppn

scales present; psp scales absent; pra scales very few, sometimes absent;

pst with a broken or complete narrow line of white scales; stp with horizontal

scale lines widely separated; vertical stp scale line not extending dorsad of

level of lower edge of mep, with or without some semierect scales at each end;

lower anterior mep scale patch narrow, far separated from upper patch. Legs:

fore- and midcoxae with upper scales white, lower usually brown, hindcoxa

with all scales dingy-white or with some lower ones brownish; trochanters,

particularly on fore- and midlegs, usually with mixed brown and white scales.

Wing: largely brown- or black-scaled; veins behind vein Rj with narrow to

medium-wide scales; scales on vein 1A wide spreading. Haltere: scales broad.

SYSTEMATICS. I believe this is the most primitive subgroup of the Sig-

nifera group. It contains the widespread kummi and species 5.

DISTRIBUTION. Arizona, south to Panama.

0. Orthopodomyia sp. Mexican highlands form

Figs. 2,11

FEMALE (fig.2). Wing: 4.52 mm. Proboscis: 2.18 mm. Forefemur:

2.32 mm. Abdomen: about 2.9mm. Head: integument of head capsule, cly-

peus, torus and flagellum dark brown; erect scales black; labium largely dark-

scaled, white scales very few, scattered; palpus about 0.45 of proboscis, large-

ly dark-scaled, segments 2 and 3 with only very few scattered white scales.

Thorax: integument of mesonotum and scutellum dark chestnut-brown, post-

notum lighter chestnut-brown, pleuron dark chestnut-brown anteriorly, lighter

posteriorly. Legs: integument of forecoxa chestnut-brown, midcoxa Somewhat

lighter, hindcoxa much lighter; legs with dark-scaling dark brown; forefemur

with anterior surface largely dark-scaled, white scales scattered along dorsal

margin near base and entire ventral margin; posterior surface with medium-

wide dark streak from base to apex, areas dorsad and ventrad of streak white-

scaled with dark speckles; midfemur with anterior surface largely dark-scaled,

_ white speckles apparently concentrated in median longitudinal streak, posterior

surface pale-scaled basally and to or near apex in progressively narrowing :

streak along ventral margin, remainder dark-scaled; hindfemur with anterior

surface white-scaled ventrally in basal portion, remainder dark-scaled with

Zavortink: Genus Orthopodomyia | ae

white speckles, dorsal margin white-scaled to about middle, posterior surface

pale-scaled basally and with white speckles ventrally near apex, remainder

dark-scaled; tibiae largely dark-scaled; foretibia with line of white speckles on

anterior surface and with at least some white speckles on posterior surface;

midtibia with white speckles on anterior, dorsal and posterior surfaces; hind-

tibia with all surfaces speckled with white scales; tarsi largely dark-scaled;

fore- and midtarsi with small dorsal white patch on at least base of segment 1;

hindtarsal segment 1 with posteriorly incomplete basal ring and with complete

apical ring, segment 5 all white-scaled dorsally. Wing: vein R with remigium

white; costa and vein Sc apparently all dark; remainder of vein R and all other

veins dark-scaled with at least a few white speckles. Haltere: integument of

stem whitish to tan, knob reddish-brown. Abdomen: integument of tergite I and

sternites II-VIII tan; abdomen with dark scales brown to blackish; tergites II-

VIII with large basolateral white patch, and at least on tergites II and III, with

middorsal white patch; sternite II entirely white-scaled; sternites IJI-VII white-

scaled basally or basolaterally, remainder dark; sternite VIII with white and

dark scales laterally.

MALE, PUPA. Unknown.

LARVA (fig.11). Head: 1.03 mm. Siphon: 0.86 mm. Anal Saddle: 0.55

mm. Not all branched hairs with conspicuous barbs. Head: largely brown col-

ored, lighter cephalad and around imaginal eye; mental plate tan, usually with

9,10 (9-11) teeth on each side; hair 2-C sometimes present; 9-C normally sin-

gle, fine; 11-C weakly developed, usually 2, 3b (2-4); 13-C moderately devel-

oped, usually 2,3b. Antenna: uniformly straw-colored. Thorax: epidermal

pigment absent or weakly developed and pink in color; spicules not conspicuous;

hairs strongly pigmented; hair 0-P usually 8, 9b (7-11); 6-P single; 1-M very

strongly developed, long, usually 3-5b (3-6); 1-T very long, usually single

(single, double); 13-T very strongly developed, long, usually 8-10b (6-11). Ab-

domen: segment VII usually with sclerotized plate and segment VI often with

plate; hairs strongly pigmented; hair 1-I moderately developed, usually 2, 3b;

4-I shorter than 3-I; 6-I usually 10, 11b (9-12); 7-I single; 1-II very long, sin-

gle; 4-II well developed; 5-II-VI moderately developed, usually 3b (2-4); 6-II

usually 12-14b (10-14); 7-II moderately developed; 1-III very long, single; 13-

IlI-V usually single or double (1-3b) and shorter than or subequal to hair 10 of

corresponding segment; 1-VI strongly developed, usually single or double (1-

3b); 1-VII very strongly developed; 13-VII usually 3,4b (3-5). Segment VIII:

large sclerotized plate present; anterior comb scales usually 23-27 (19-32),

posterior comb scales usually 6-9 (5-12); posterior row of comb scales about

0.43-0.53 length of anterior row; hair 3-VIII very strongly developed, usually

8-10b (7-10). Siphon: index about 2.6-3.3; uniformly brown in color; integu-

mentary sculpturing absent; hair 1-S located 0.50-0.54 from base of siphon,

9-11b, very strongly developed and with longest branches extending far beyond

apex of siphon. Anal Segment: saddle complete, light brown; integumentary

sculpturing inconspicuous, imbricate; diameter of saddle at most proximal

point not or only slightly smaller than diameter at the most distal point along

ventral surface, so that dorsal and ventral surfaces are parallel or diverge on-

ly slightly when viewed from the side and diameter at most proximal point of

saddle usually equal to or greater than diameter of base of siphon; hair 1-X

single, fine; ventral brush with 7 pairs of hairs; more strongly developed hairs

in ventral brush with 8-10 branches; gills long, dorsal pair about twice as long

as ventral pair and 1. 4-2.4 length of anal saddle.

- BIONOMICS. Larvae have been collected in a large treehole at ground lev-

el. Nothing is known of the habits of the adult.

SYSTEMATICS. This species is known from a single badly rubbed female

60 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

collected at 9,000 feet elevation at El Salto, Mexico. It is quite similar to kum-

mi in most details, but differs from it in having the line of white scales on vein

R restricted to the remigium and fewer white speckles on the proboscis and

palpus than all but the depauperate specimens of kummi.

I am tentatively associating larvae collected at about 8, 400 feet elevation

near Zitacuaro (about 450 miles south of EF] Salto) with this adult. These larvae

are extremely hairy forms; they resemble, but are even more hairy than, the

hairy forms of kummi from Guatemala and signifera from Arizona and Califor-

nia. In particular hairs 13-T and 6-II are more strongly developed in these

larvae than in the hairy forms of kummi and signifera.

While I believe that the adult and larva are the stages of a distinct Species

from the Mexican highlands, I am not naming the adult because it is known from

only one damaged specimen and am not naming the larva because of the possi-

bility that it is only a hairy form of kummi.

DISTRIBUTION. Known only from the Mexican states of Durango and Mex-

ico. Material Examined: 70 specimens; 1 9, 69 L.

MEXICO. Durango: El Salto (10 miles W), 10 July 1964, J. F. McAlpine,

1 2? [CNC].

Seiad ae Zitacuaro (28 miles E), 3 Sept 1964, E. Fisher and D. Verity »

(MEX 167), 69 L [UCLA].

6. Orthopodomyia kummi Edwards

Figs. 2,12,13

1939. Orthopodomyia kummi Edwards, 1939:121-122. *TYPE: holotype °,

Orosi [Cartago|, Costa Rica, reared from a larva found in an abandoned

tank or cesspool full of clear rainwater, 16 Dec 1937, H.W. Kumm

[BMNH].

Orthopodomyia kummi of de Buen (de Biagi) (1952:243-252; 1953:189, 191); Lane

(1953:626); Vargas (1956:31); Stone, Knight and Starck (1959: 123): McDon-

ald and Belkin (1960:249-250); Storie (1961:36); Diaz Najera (1963:191); Bur-

ger (1965:396); Chapman (1965:432, 435, 436, 438); Stone, Sabrosky, Wirth,

Foote and Coulson (1965:108); Diaz Najera (1966: 63).

Orthopodomyia n.sp. of Kumm, Komp and Ruiz (1940:405, 417, 418).

Orthopodomyia signifera of Richards, Nielsen and Rees (1956: 16): McDonald

(1957a:505; 1957b:535).

Orthopodomyia californica of Rigby and Ayres (1961:56).

FEMALE (fig. 2). Wing: 4.65 mm. Proboscis: 2.82 mm. Forefemur:

2.82 mm. Abdomen: at 3.1mm. Head: integument of head capsule, cly-

peus, torus and flagellum reddish-brown to black; erect scales brown to black;

labium largely dark-scaled, white scales few to numerous, scattered, yet

somewhat concentrated along dorsal surface; palpus about 0.41 of proboscis,

largely dark-scaled, segments 1,2 and 3 usually with some scattered white

scales. Thorax: integument of mesonotum, scutellum, and postnotum evenly

reddish-brown to black, pleuron more or less evenly reddish-brown to black

anteriorly and dorsally, lighter on central posterior portion of stp, lower por-

tion of mep and on metameron. Legs: integument of forecoxa reddish-brown

to dark brown, midcoxa somewhat lighter, hindcoxa much lighter; legs with

dark scales brown to black; forefemur with anterior surface usually streaked

with white scales along dorsal margin basally, remainder dark-scaled with

Zavortink: Genus Orthopodomyia 61

white scales scattered along apical dorsal margin and entire ventral margin,

posterior surface with narrow to broad, median or more ventral, dark streak

from base to apex, areas dorsad and ventrad of streak entirely white-scaled or

with dark speckles; midfemur with anterior surface usually largely dark-scaled

with white speckles dorsally, posterior surface pale-scaled basally and to apex

in progressively narrowing streak along ventral margin, remainder usually

dark-scaled with white speckles, sometimes all dark-scaled; hindfemur with

anterior surface usually white- or pale-scaled ventrally in basal portion, some-

times narrowly white-scaled across entire base, remainder largely dark-

scaled with few to many white speckles, dorsal margin usually white-scaled to

beyond middle, posterior surface more or less like that of midfemur, often

with basal pale-scaling extending farther apically; tibiae largely dark-scaled;

foretibial anterior surface usually with white scales in line or line of speckles

from base to or near apex, posterior surface with similar but shorter line or

with very sparse or irregular white speckling; midtibial anterior and posterior

surfaces with more or less distinct white lines from base to or near apex, ven-

tral and sometimes dorsal surfaces speckled with white; hindtibial anterior,

posterior and usually dorsal surfaces speckled with few to numerous white

scales, which, when numerous, may form more or less distinct lines basally

on anterior and posterior surfaces; tarsi largely dark-scaled; foretarsal seg-

ment 1 usually with very small to small dorsal white patch at base; midtarsal

segment 1 with medium-sized dorsal white patch at base and sometimes with

smaller one at apex, segments 2, 2-3 or 2-4 sometimes with a few white scales

at base; hindtarsal segment 1 with posteriorly incomplete basal white ring, and

sometimes with apical white ring, segment 5 all white-scaled dorsally. Wing:

vein R with line of white scales from base to separation of vein R,; additional

light-scaling variable within 1 population and also apparently varying clinally,

southern populations having fewer white scales, northern populations more; at

minimal development only a few scattered white scales occur on distal portion

of vein Sc and middle and distal portions of vein Rj; at maximal development

numerous scattered white scales occur on distal portion of vein Sc, middle and

distal portions of vein Rj, distal portion of stem of vein Cu, basal portion of

vein Cuy and distal portion of vein 1A and a few scattered white scales occur

on the costa and basal portion of vein R445; remaining portion of veins men-

tioned and all other veins entirely dark-scaled; scales narrow in southern pop-

ulations, wider in northern ones. Haltere: integument of stem whitish to brown,

knob tan to brown. Abdomen: integument of tergite I tan to brown, sternites II-

VII whitish to brown, sternite VIII brown to dark brown; abdominal ornamenta-

tion quite variable; abdomen with dark scales brown to black; tergite II with

white scales in large middorsal patch which may extend to apical margin and

in small to large basolateral patch or narrow to broad basal band; more prox-

imal of tergites III-VII with white scales in small to large basolateral patch,

dorsal surface all dark, with only a few white scales basally, or with conspic-

uous basal white patch; more distal of tergites III-VII with white-scaling more

extensive, basolateral white patch larger, dorsal surface more often with basal

white patch and sometimes with complete basal white band; tergite VIII varying

from largely dark-scaled to having complete narrow basal white band; sternite

II entirely white-scaled or with some scales dark; sternites III-VII with white

scales basally or basolaterally, remainder dark-scaled, more distal of these

sternites more extensively dark-scaled than proximal ones, sternite VIII with

dark or dark and white scales laterally.

MALE. Essentially as infemale. Labium: largely dark-scaled, white

scales in narrow ring just distad of level of distal end of palpal segment 2 and

62 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

scattered, especially dorsally and/or distally. Palpus: about 0.96 of probos-

cis, Segment 3 without well developed bristles; white scales scattered on seg-

ments 1,2 and 3, and sometimes concentrated near base of segment 3. Anten-

na: flagellar segment 1 with black and white scales dorsomesally. Thorax: up-

per mep bristles reduced in number. Wing: usually with fewer white speckles

than in females from the same collection, and showing, but less conspicuously,

the same clinal variation as females; more southern populations with only a

few scattered white scales on distal portion of vein Sc and sometimes on middle

and distal portions of vein Ry; more northern populations with numerous scat-

tered white scales on distal portion of vein Sc and on middle and distal portions

of vein Ry, and with vein 1A, middle portion of vein Cu, and base of vein R4+5

usually with at least 1 white scale. Abdomen: ornamentation variable, but with

white-scaling generally more extensive than in females; tergites III-VII as in

female or with narrow to broad basal white band which may be wider laterally

and middorsally, more distal of these segments usually more extensively

white-scaled than proximal ones; tergite VIII usually with white basal band;

sternites III-VII with white basolateral patch or with narrow to broad basal

white band which may be wider laterally, more proximal of these segments

usually more extensively white-scaled than distal ones; sternite VIII varying

from entirely dark-scaled to entirely white-scaled.

MALE GENITALIA (fig. 12). Segment VIII: tergite lobe broadly truncate

or emarginate, sometimes with small serrations near apex, sides parallel or

diverging apically. Segment IX: tergite with 2-6 bristles on each side; sternite

rarely with small projection in middle of cephalic edge. Sidepiece: basal mesal

lobe with 3-5 stout bristles which are bent mesally near apex and with 1-3 finer

bristles. Clasper: usually constricted distad of origin of apodeme, so that mid-

dle portion is narrower than distal portion. Aedeagus: basal portion usually

more angular than in other species; ventral triangular spines variable in posi-

tion; preapical ridge with 2 or 3 processes. Proctiger: paraproct sclerotiza-

tion usually with 2-5 apical teeth; cercal setae usually 2-5.

PUPA (fig.12). Abdomen: 3.7mm. Trumpet: 0.5mm. Paddle: 0.7 mm.

Cephalothorax: hairs 8,9-C often as long as trumpet. Trumpet: deep tan to |

brown except at extreme base and apex and largely darker than darkest part of

cephalothoracic integument. Metanotum: hair 12-C usually strongly developed.

Abdomen: hairs 6, 7-I usually strongly developed, long; 1-II usually 2, 3b (1-4),

very strongly developed, long; 2-II-VI not more than 0.1-0.3 length of hair 1 of

corresponding segment; 6-II long and strong, considerably stronger than 7-II;

O-III usually 3-5b and not nearly as strongly developed as 5-IV-VI or 3-III; 10-

III- VII usually reaching to or nearly to caudal margin of following segment; 5-

IV, V usually extending to or nearly to caudal margin of following segment.

Paddle: usually narrowly to broadly elliptic; apex rounded, truncate or emar-

ginate; inner part usually with band of minute spicules near inner margin.

LARVA (fig.13). Head: 0.91 mm. Siphon: 0.72 mm. Anal Saddle: 0.43 .

mm. Not all branched hairs with conspicuous barbs. Head: largely light brown

to brown in color, lighter cephalad and around imaginal eye, collar darker;

mental plate brown, usually with 9,10 (8-11) teeth on each side; hair 2-C ap-

parently absent; 9-C single or double, fine; 11-C weakly developed, usually

with 3, 4b (3-5); 13-C moderately developed, usually 2, 3b (2-4). Antenna: uni-

formly light tan colored. Thorax: epidermal pigment weakly to strongly devel-

oped, pink to red or purple in color; spicules not conspicuous; hairs strongly

pigmented; hair 0-P usually 5, 6b (4-8); 6-P single; 1-M usually moderately

developed and single, strongly developed and 2, 3b in hairy form; 1-T usually

weakly developed and single, strongly developed and 1-3b in hairy form; 13-T

Zavortink: Genus Orthopodomyia : 63

usually 3-5b and moderately developed or, especially in hairy form, strongly

developed. Abdomen: segment VII usually with sclerotized plate and segment

VI often with plate; hairs strongly pigmented; hair 1-I usually weakly developed

and 2, 3b (1-4), strongly developed and 1-3b in hairy form; 4-I shorter than 3-];

6-I usually 7, 8b (6-9); 7-I single; 1-II usually weakly developed and single (sin-

gle, double), strongly developed and single or double in hairy form; 4-II weakly

to moderately developed; 5-II-VI usually weakly to moderately developed and

2,3b (2-4), quite strongly developed in hairy form; 6-II usually 6-8b; 7-II usu-

ally moderately developed, strongly developed in hairy form; 1-III usually mod-

erately developed and single (single, double), strongly developed and single or

double in hairy form; 13-III-V usually single or double (1-3b) and shorter than

hair 10 of corresponding segment; 1-VI usually weakly developed and double

(1-3b), moderately developed and double (single, double) in hairy form; 1-VII

usually moderately developed and double (single, double), strongly developed

and 2, 3b in hairy form; 13-VII usually 4, 5b (4-6). Segment VIII: large sclero-

tized plate present; anterior comb scales usually 18-22 (17-26), posterior comb

scales usually 5-7 (5-9); posterior row of comb scales about 0.33-0.58 length

of anterior row; hair 3-VIII strongly developed, usually 7-9b (7-11). Siphon:

index about 2.7-3.8, usually less than 3.5; largely light brown to brown in col-

or, extreme base darker, apex sometimes lighter; integumentary sculpturing

absent to weakly developed, imbricate; hair 1-S located 0.48-0.57 from base

of siphon, usually 9-11b (8-12), strongly developed and with longest branches

extending considerably beyond apex of siphon. Anal Segment: saddle complete;

saddle uniformly light brown to brown or somewhat darker dorsoapically; in-

tegumentary sculpturing weakly developed, imbricate; diameter of saddle at

most proximal point usually considerably smaller than diameter at the most

distal point along the ventral margin, so that dorsal and ventral surfaces di-

verge when viewed from the side, and diameter of saddle at most proximal point

usually less than diameter of base of siphon; hair 1-X usually single and fine,

often enlarged and branched in hairy form; ventral brush usually with 7 pairs

of hairs; more strongly developed hairs in ventral brush usually with 11-13

branches (9-17); gills usually long, dorsal pair about 1.4-1.7 as long as ven-

tral pair and 1.1-2.1 length of anal saddle.

BIONOMICS. Larvae of kummi have been found in treeholes, open-ended

bamboo internodes, artificial containers and in an abandoned cesspool. They

are found in the same treeholes as signifera in southeastern Arizona. Nothing

is known of the habits of adults except that they are active during darkness and

will take avian blood (see Biology and Ecology section).

SYSTEMATICS. Kummi is similar to species 5 but is told from it and all

other species in the Signifera group in the adult stage by having the line of white

scales on the base of vein R extending to the separation of Rg.

Adults of this species are quite variable. Wing variation is clinal, with

northern populations having broader and more white scales than southern popu-

lations. This cline is evidently due to introgression with signifera, which oc-

curs with kummi in southeastern Arizona and is known to hybridize with it. The

description of the wing of northern populations is based on individuals collected

in areas of Arizona where signifera does not exist. In those areas where the 2

Species occur together, there are some females which appear to be kummi on

the basis of all larval, pupal and adult features except that they have a few ad-

ditional white scales on any or all of the following veins: base of veins Rg,

R9+3, Rg, Cu and 1A and on vein M near the crossveins. I have considered

these individuals to be kummi, but they may actually be backcrosses of kummi

to the Fj hybrids between kummi and signifera. The hybrids are discussed be-

64 Contrib. Amer. Ent. Inst., vol. 3, no. p. 1968

low. Femoral ornamentation in collections from treeholes separated by only a

few meters often differs considerably, leading me to believe that the differences

are environmentally induced rather than having a genetic basis. Other varia-

tions in adult ornamentation are due to larval nutrition; depauperate adults have

the white-scaling on the abdomen, palpus, proboscis, femora and tibiae much

reduced. | |

While most larvae throughout the species range are non-hairy, extremely

hairy individuals have been found in Guatemala and partially hairy larvae occur

at Patagonia, Arizona. The non-hairy larvae are difficult or impossible to dis-

tinguish from signifera from the eastern United States and the Greater Antilles,

and from pulchripalpis and waverleyi. The hairy larval form is similar to spe-

cies 5 and the hairy form of signifera. Except for a single specimen collected

in bamboo in El Salvador, the pupa of kummi is the hairy form and is similar

to that of signifera from the eastern United States. It is generally told from the

latter by the greater development of hairs 6, 7-I, 1,6-II and 5-IV-VI and espe-

cially by the development of minute spicules near the inner margin of the pos-

terior portion of the paddle. These spicules are present in all or most individ-

uals of every collection of kummi, but are very rarely present in signifera and

never occur in any other species of the Signifera group.

Kummi and signifera occur together at some localities in southeastern Ari-

zona and, to date, specimens appearing to be hybrids between the species have

been found at the following three locations in Cochise County: Huachuca Moun-

tains, Coronado National Memorial; Dragoon Mountains, Cochise Stronghold:

Campground; and Chiricahua Mountains, Chiricahua National Monument. Sup-

posed hybrid adults are similar to kummi except that the pleural scale lines

are broader and the wings have numerous scattered white scales and may even

. have a weakly developed spot over the crossveins. The number of scattered

white scales is generally greater than that indicated above for the possible

backcrosses to kummi. Pupae of the hybrids are nearly indistinguishable from

those of kummi, but do have some hairs, particularly 1-II and 4-V, less well

developed than that species. Because of variation present in the parental spe-

cies, hybrid larvae are difficult to distinguish. In southeastern Arizona, larvae

of kummi generally differ from those of signifera by the following features: (1)

hair 1-S with a greater number of branches and the branches more evenly

spaced, (2) hair 1-T,II shorter and more weakly developed, (3) siphon more

uniformly sclerotized and with straighter lines, (4) epidermal pigment more

frequently present and darker in color when present, (5) abdominal plates larger

and more numerous, (6) anal gills longer, and (7) sclerotized parts darker

in color. Hybrid larvae usually resemble kummi in character 2 and sometimes

in character 6, but, for the most part, tend toward signifera in the other char-

acters. They normally differ from the larvae of either pure species by having

a longer siphon. However, it cannot be overemphasized that the larval varia-

tions of each parental species is so great that they sometimes cannot be told.

from each other and, as a consequence, it is difficult to recognize the hybrids.

I am considering kummi to be the most primitive species in the Signifera

group because the adult shares many features with albicosta, a South American

species which I have placed in a separate group. The wing and pleural orna-

mentation of the 2 species are almost identical; in addition, kummi tends to-

ward albicosta by having a more triangular clypeus, fewer scales between the

eyes and fewer pra bristles than any other species in the Signifera group. It is

of course possible that these similarities are due to introgression rather than

relationship.

Zavortink: Genus Orthopodomyia 65

Kummi

DISTRIBUTION. Southeastern Arizona, south to northern Panama. Ma-

terial Examined: 2925 specimens; 693 o, 5859, 3 A, 801 P, 843L; 286 indi-

vidual rearings (213 larval, 47 pupal, 26 incomplete).

COSTA RICA. Orosi, 16 Dec 1937, H.W. Kumm, 26 o, 38 2, 10L [USNM]

holotype 2, 40, 3 2? [BMNH], 1, 1 A [UCLA].

EL SALVADOR. Santa Tecla, 19 June 1958, O.L. Cartwright, 1L [USNM].

Sonsonate, Canton El Castano, 1 Aug 1964 (SAL 2), 1 po (2-100) [UCLA].

GUATEMALA. Solola, 30 June 1965, J.N. Belkin and T.J. Zavortink

(GUA 25), 1 lpo (25-12), 3 po (25-100-102), 4 1p (25-10, 11,13,14), 1P, 54L

[UCLA]. No locality or date (GUA 151), 10, 29, 2 P [UCLA].

MEXICO. Baja California Sur: Todos Santos, 30 May 1965, W.A. McDon-

ald (UCLA 237), 201po (237-11-29, 31), 21p92 (237-10,30), 1 po (237-102),

4 p? (237-101, 103-105), 13%, 2 9, 16 P, 37 L [UCLA]; 1 June 1965, W.A.

McDonald (UCLA 246), 3 lpo (246-11, 12,14), 1 1p? (246-10), 5 po (246-102-

106); 2.p¥ (286-100, 101), Toto (240-12) 1 21 PS oe CA),

Chiapas: Mapastepec, El Pavat, 20-27 July 1959, B. Lievano, 1 o, 3 L

[ISET].

Guerrero: Omilteme, 8 Sept 1965, D. Schroeder (MEX 361), 14L [UCLA].

Oaxaca: Las Cruces, El Barrio, 16 Oct 1959, H. Calderon, 10°, 2 9 [ISET].

San Luis Potosi: Tamazunchale, 20 July 1965, R.X. Schick and D. Schroeder

(MEX 233), 1 1po (233-20) [UCLA]. Tlapexhuncan, May 1942, 10%, 1 9 [USNM ].

Veracruz: Cordoba, 22 July 1964, E. Fisher and D. Verity (MEX 71),

Llp? (71-10), 20%, 49, 4L [UCLA]; 4 Aug 1964, E. Fisher and D. Verity

(MEX 93), 2 1p (93-11,12), 1 L [UCLA].

PANAMA. Chiriqui: El Volcan Chiriqui, 6 Mar 1943, T.H.G. Aitken, 1 0

[UCLA]; 30 June 1943, T.H.G. Aitken, 19, 40 L [USNM], 2 2 [UCLA].

UNITED STATES. Arizona: Chiricahua National Monument, 24 Dec 1966,

L. T. Nielsen (N-37-66), 6%, 19, 1 A[UTAH]. Coronado National Memorial,

25 Dec 1966, L.T. Nielsen (N-41-66), 20°, 29, 3 P, 4L [UTAH]; same data

(N-42-66), 140°, 69, 4 P [UTAH]. Madera Canyon, 22 Aug 1954, W.A. Mc

Donald (UCLA 135), 21p (135-101,111), 3 L [UCLA]; 17-19 Aug.1955, W.A.

McDonald (UCLA 171), 15 lpo& (171-303, 304, 306, 307, 309-315, 325, 326, 328,

349), 26 lp (171-316-320, 323, 324, 327, 329-332, 334-343, 345-348), 5 po” (171-

301, 302, 322, 351, 352), 1 1o° (171-321), 3 P, 24 L [UCLA], 2 1po (171-305,

308) [USNM]; 26 Dec 1966, L.T. Nielsen (N-43-66), 210°, 49, 1A, 8P,

6L[UTAH]; Nogales, 21 Mar 1966, T.J. Zavortink (UCLA 298), 3 lpo (298-

10,17,19), 71p@ (298-11-16, 18), 240, 289, 52 P, 40 L [UCLA]; same data

(UCLA 299), 6 1po (299-10, 11, 13-16), 2 lp? (299-12,17), 20, 2P, 2L [UCLA].

Patagonia, 24 Aug 1954, W.A. McDonald (UCLA 137), 7L [UCLA]; 18 Aug

1964, J. Burger (UCLA 252), 3 po’ (252-101, 102, 104), 1 p? (252-103), 91 ©,

1179, 1P [UCLA]; same data (UCLA 253), 2 po’ (253-104, 106), 4 p? (253-101-

103,107), 92, 999, 4 P [UCLA]; 13 Sept 1964, J. Burger (UCLA 259), 24,

16 2 [UCLA]; 24 Jan 1965, J. Burger (UCLA 273), 107 ¢, $8 9, 30 BY 71-L

[UCLA]; 25 July 1965, J. Burger (UCLA 280), 3 L [UCLA]; 27 July 1965, J.

Burger (UCLA 282), 26 L [UCLA]; 21 Mar 1966, T.J. Zavortink (UCLA 300),

10 1lpo& (300-10-14, 16, 19, 28, 31,37), 11 1p (300-15, 17, 18, 20-25, 27, 29), 11p

(300-26), 26, 112, 39 P, 78 L [UCLA]; 5 Sept 1966, T.J. Zavortink (UCLA

333), 6 lp? (333-14-19), 50°, 39, 9 P, 19 L [UCLA]; same data (UCLA 334),

a ipo (saa-11, 21-23), 4 dps. dee 7 10,2100, 24.85), Jac. 7 26 BG 1,

[UCLA]; same data (UCLA 335), 2 1po (335-11, 12), 3 1p? (335-10, 13,14), 1 p?

(335-100), 50%, 6 9, 11 P, 7L [UCLA]. Pearce, Cochise Stronghold Camp-

66 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

ground, 26 Aug 1964, J. Burger (UCLA 255), 6 po (255-122, 127-131), 6 p

(255-120, 121, 123-126), 150’, 32, 1 P [UCLA]; 22 Mar 1966, T.J. Zavortink

(UCLA 802), Gipe (3402-18-20 22-24), 1 1p° 602-21), a0, 10°. 90 P, 41.

[UCLA]; 4 Sept 1966, T.J. Zavortink (UCLA 325), 2 lp? (325-10, 11), 2P

[UCLA]; same data (UCLA 326), 21 1p? (326-11-17, 20, 22-25, 27-32, 34-36),

5 1p (326-10, 18, 21, 26, 33), 62, 9P, 1L [UCLA]; same data (UCLA 328), 4 1po'

(328-31, 32, 35,37), 10°, 1P, 1L [UCLA]; same data (UCLA 329), 16 1p? (329-

20,21, 23,25, 26, 26-30, 92-39), 41 (820-22 94.97.91), 46-6 920, 2490, 67 L

[UCLA]; 6 Sept 1966, T.J. Zavortink (UCLA 342), 3 lpo (342-12, 14, 27), 5 1p

(342-16, 18-21),.1 Lp-(842-11),.10.¢,.12 P, 21, [UCLA]. Sierra Vista, Carr

Canyon, 5 Sept 1966, T.J. Zavortink (UCLA 338), 2 lpo (338-11,12), 6 lp?

(38-13-16, 18,19), 1 1p (338-17), 20°, 2 P, 6L [UCLA]; same data (UCLA

340), 2 1p? (340-14, 15), 2 po’ (340-102, 103), 4 1p (340-10-13), 2, 9P [UCLA];

‘game data (UCLA 341), 3 lpo (341-10, 13,14), 5 1p? (341-12, 15-18), 1 po"

(341-100), 10%, 2 P, 4L [UCLA]. Sierra Vista, San Pedro River, 5 Sept 1966,

T.J. Zavortink (UCLA 336), 22 [UCLA]; same data (UCLA 337), 1 lpo (337-

17), 3 1p? (337-10, 18,19), 3.0)..2 005 P, 8. WCLAL

Additional records from the literature. MEXICO. Puebla: El Agengibre,

31 May-19 July 1952, D. Garza, F. Biagi and A.M. de Buen, L, P (de Buen

(de Biagi), 1952:243-252).

Hybrids

DISTRIBUTION. Known from only Cochise County, Arizona. Material

Examined: 69 specimens; 40, 149, 9 P, 42 L; 9 larval individual rearings.

UNITED STATES. Arizona: Chiricahua National Monument, 24 Dec 1966,

L.T. Nielsen (N-37-66), 22 [UTAH]. Coronado National Memorial, 25 Dec

1966, L. T. Nielsen (N-42-66), 59, 6 L [UTAH]. Pearce, Cochise Stronghold

Campground, 22 Mar 1966, T.J. Zavortink (UCLA 302), 2 lpo& (302-10, 11),

4 1p? (302-12-15), 22, 7 L [UCLA]; 4 Sept 1966, T.J. Zavortink (UCLA 326),

1 lpo (826-37), 1 L [UCLA]; 6 Sept 1966, T.J. Zavortink (UCLA 342), 1lpo

(342-26), 1 lp? (342-29), 19 L [UCLA].

Additional records from the literature. None.

ALBICOSTA GROUP

7. Orthopodomyia albicosta (Lutz)

Figs. 2,14,15

1904. Bancroftia albicosta Lutz, 1904:6,1. TYPE: A, [Serra da Cantareira,

S40 Paulo], Brazil [Location unknown]. Neither of the specimens in the

BMNH marked as types by J. Lane can be part of the type series because

of different locality (Cochocirinha) or date (1905) later than that of the

original description.

Orthopodomyia albicosta of Dyar (1928:398); Edwards (1932:108); Prado (1935:

1510); Edwards (1939:121, 122); Lane (1939:96); Anduze (1941:11); Davis

(1944:230); Lane (1953:624-625); Stone (1957:334); Stone, Knight and Starcke

(1959:122); Cova-Garcia, Sutil and Rausseo (1966a:53; 1966b:51, 117, 370-

371).

Orthopodomyia (Bancroftia) albicosta of Lima (1935:175-176).

Bancroftia albicosta of Blanchard (1905:632); Lutz (1905:67-69); Theobald (1907:

521-523); Peryassu (1908:21, 70, 247-249, 354); Theobald (1910b:469); Peryas-

Zavortink: Genus Orthopodomyia 67

su (1929:281).

FEMALE (fig. 2). Wing: 4.15 mm. Proboscis: 2.32 mm. Forefemur:

2.38 mm. Abdomen: about 2.7mm. Head: integument of head capsule and

flagellum brown, clypeus tan, torus dark brown mesally, beige to tan laterally;

eyes contiguous above antennae; orbital bristles usually 1 lateral pair, 2 mesal

pairs; vertex and occiput with numerous decumbent scales, those on vertex

black, broad and flat, those on occiput white, broad and flat laterally, narrow and

curved mesally, erect scales brown to black; dorsolateral scales all white or

some black; lateral and ventral scales white; suborbital bristles tan or brown;

clypeus triangular, bare; labium largely dark-scaled, white scales scattered

over entire surface or more or less concentrated dorsally, and in short mid-

dorsal longitudinal streak in apical half; palpus about 0.46 of proboscis, 4-seg-

mented, segment 4 very small and entirely silvery-white-scaled, segments 1-3

largely dark-scaled, with scattered white scales on segments 2 and 3; torus

with mesal line of small silvery-white flat scales; flagellum with segment 1

slightly swollen, no segments with scales; dorsal flagellar bristles about 3.0

times length of their segment on proximal segments; ventral bristles absent on

segment 1 and reduced in length on segments 2-4. Thorax: integument of mes-

onotum deep reddish-brown to brownish-black at margins, along dorsocentral

bristles and in posterior quarter, remainder lighter, either yellowish-tan, light

brown or sometimes reddish-brown, scutellum brown to blackish-brown, post-

notum reddish- to blackish-brown, usually with darker median longitudinal

streak, pleuron brown except for lighter areas near sutures and posterior por-

tion of mep; paratergite moderately broad; humeral and lateral prescutal bris-

tles few, posterior fossal bristles absent; mesonotal ornamentation consisting

of 4 pairs of lines of small white scales arranged into a pattern of 2 pairs of |

longitudinal lines, and broader bands of small brown scales; white scales ar-

ranged as follows: (1) lateral prescutal scales in line from anterior promon-

tory to scutal suture laterad of humeral and lateral prescutal bristles, (2) su-

praalar scales in line from scutal suture to parascutellum mesad of supraalar

bristles, (3) outer dorsocentral scales in line from caudad of humeral bristles

to lateral prescutellar scale line laterad of dorsocentral bristles, mesad of

fossal bristles, (4) lateral prescutellar scales much larger and longer than

other mesonotal scales, in line from anterior edge of prescutellar space to

scutellum mesad of lateral prescutellar bristles, connected anteriorly to outer

dorsocentral scale line; brown scales arranged as follows: (1) in band accom-

panying, but extending further caudad than, acrostichal bristles, (2) in band

accompanying dorsocentral bristles; lateral lobes of scutellum with or without

a few long, thin, white scales, midlobe with 1 pair of narrow white scale lines

which extend 3 to 4 times length of scutellum from its caudal margin; upper

anterior apn bristles 1-4 strongly developed, 1-4 weakly developed, lower an-

terior apn bristles 1 or 2, moderately to weakly developed, tan; ppl bristles

4-6 strongly developed, 2-5 weakly developed; pra bristles absent; upper stp

bristles 1, strongly developed, posterior stp bristles 2 strongly developed, 0

or 1 weakly developed, lower stp bristles 1 or 2, weakly to moderately devel-

oped, amber in color; upper posterior mep bristles 4-7, amber, anterior mep

bristles absent; pleural scaling with individual lines narrow; apn with diagonal

row or curved decumbent scales between upper and lower bristles; ppn with

horizontal line of curved decumbent scales extending from line on apn to its

caudal border ventrad of posterior ppn bristles; psp bare; pra with many curved

decumbent scales; ppl with row of curved, decumbent scales below ppl bristles;

pst with curved decumbent scales near ppl; pcx, ssp and hypostigial scales ab-

68 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

sent; stp with 3 distinct lines of scales, as follows, decumbent scales in 2 hor-

izontal rows, 1 above and 1 below upper stp bristle, and broad decumbent and

semierect scales in nearly vertical line extending ventrad from level of lower

margin of mep along posterior stp margin; mep with upper horizontal line of

decumbent and long semierect scales and lower horizontal line of decumbent

scales; additional large brownish scales may be scattered on pleuron. Legs:

integument of forecoxa tan in upper portion, remainder beige, midcoxa brown

in upper outer portion, remainder beige, hindcoxa beige; fore- and hindcoxal

bristles beige to tan, midcoxal bristles brown; broad to rather narrow, flat

scales present along outer edge of anterior surface on all coxae and above and

behind uppermost bristles on forecoxa, coxae largely white-scaled, light brown

scales usually above bristles and along anterior surface of forecoxa and on low-

er portion of hindcoxa; trochanters with narrow, appressed scales, those on

anterior and mesal surfaces light brown, those on posterior surface white; re- _

maining leg segments with dark scales brown to black; femora without knee

spots; forefemur with anterior surface white-scaled at base, remainder dark-

scaled with white speckles, the latter often more common apically, posterior

surface irregularly but almost completely white-scaled ventrally, dark-scaled

with white speckles dorsally; midfemur with anterior surface largely dark-

scaled, white scales in median longitudinal line from near base to apex, and

scattered, especially dorsally, posterior surface white-scaled basally and to

or near apex in progressively narrowing longitudinal streak at or near ventral

margin, remainder dark-scaled with white speckles; hindfemur with anterior

surface white-scaled in ventral longitudinal streak which narrows and continues

to apex as median line, white scales scattered along dorsal margin, remainder

dark-scaled, posterior surface largely white-scaled, dark scales present dor-

sally in distal portion and along ventral margin near apex; tibial largely dark-

scaled, without apical white patches; foretibial anterior surface with white

scales in poorly defined line in basal portion or in well defined line to near

apex, posterior surface with irregular light mottles or speckles in poorly de-

fined line in basal portion or in well defined line to near apex; midtibial ante-

rior and posterior surfaces with well defined white line from base to or near

apex; hindtibial anterior surface with well defined white line from base to apex,

posterior surface with similar line in basal 0.33 and streak of tannish scales

from there to near apex; tarsi largely dark-scaled; foretarsus entirely dark-

scaled or with a few dorsal white scales at base of segment 1 and over joint be-

tween segments I and 2; midtarsus with dorsal white patch at base of segments

1-3 or 4 and apex of segments 1-2,3 or 4, basal patch usually larger than api-

cal; hindtarsal segment 1 longer than hindtibia; hindtarsus with dorsal light

patch at base of segments 1-4 or 5 and apex of segments 1-3 or 4, patch at base

of segment 1 small, patch over joint between segments 1 and 2 large, usually _

forming narrow ring ventrally, patch at apex of segment 4 or base of segment

0 very Small when present. Wing: all veins entirely dark brown-scaled except —

for vein R, which is white-scaled from base to separation of vein Rg; veins

posterior to vein Rj with narrow scales; scales on vein 1A only slightly spread-

ing. Haltere: integument of stem whitish to beige, knob brown; scales on stem

white and brown, those on knob all brown. Abdomen: integument of tergite I

beige to tan, sternites II-VII whitish to brown, apex of sternites often darker

than base and more distal sternites darker than proximal, sternite VIII brown;

tergite I fan brown- or brown- and white-scaled; remaining abdominal segments

with light scales white or creamish, dark scales brownish-black; tergite II with

large middorsal basal light patch and basolateral light patch, remainder dark-

scaled; tergites III-VII with basolateral light patch, remainder dark-scaled;

Zavortink: Genus Orthopodomyia 69

tergite VIII like preceding or with additional light scales basally; sternite II

and sometimes III entirely light-scaled; sternites IJI-VII usually white-scaled

basally, dark-scaled apically or dorsoapically, dark-scaling more prevalent

on distal segments; sternite VII mostly bare midventrally, dark and light scales

present laterally.

FEMALE GENITALIA. Tergite IX without setae.

MALE. Essentially as infemale. Labium: largely dark brown- to black-

scaled, white scales forming short longitudinal streak on dorsal surface near

apex, ventral patch distad of level of distal end of palpal segment 2 and speck-

led over surface, especially distally. Palpus: about 0.77 of proboscis, seg-

ment 5 very short, segment 3 without well developed apical bristles; white |

scales scattered on segments 2 and 3, in patch at base of segment 4 and on en-

tire surface of segment 5. Antenna: integument of flagellar segments 1-11

whitish, that of segments 12 and 13 dark brown; segment 1 of flagellum without

scales; bristles of flagellar whorls about 0.45 length of entire flagellum on prox-

imal segments, about 0.6 to 0.9 length of segment 13 on that segment. Abdo-

men: tergite VIII mostly light-scaled, sternite VIII dark-scaled with light baso-

lateral patch. : ,

MALE GENITALIA (fig. 14). Segment VIII: tergite lobe distinct, rather

Small, narrowing apically, apex rounded. Segment IX: tergite without bristles.

Sidepiece: medium-long conical; basal mesal lobe with 4 or 5 strong terete

bristles and 1 or 2 finer sternally; mesal surface membranous for short dis-

tance distad of basal mesal lobe; mesal surface with specialized bristles in

large area distad of basal mesal lobe, individual bristles directed tergomesal-

ly, with ends fine and curved sternally; specialized tergomesal bristles absent.

Spiniforms: 2, simple. Aedeagus: strongly sclerotized; apex broadly truncate,

with row of 6-8 crenations on each side, without ventral spines or processes,

but with small basal projection extending cephalad between ventral parameres.

Proctiger: paraproct sclerotization with 2 or 3 apical teeth; cercal setae 3-5.

PUPA (fig. 14). Abdomen: 3.0 mm. Trumpet: 0.4mm. Paddle: 0.6 mm.

Cephalothorax: lightly to moderately pigmented, with dorsal portion of wing case

and portion of mesonotum caudad of trumpet usually darker; integumentary

sculpturing absent; hairs concolorous with to slightly darker than integument;

all hairs shorter than trumpet; hairs 1,2-C subequal, moderately developed,

usually single or double (1-3b); 3-C weakly developed, much shorter than 1 or

2-C; 4,5-C subequal, moderately developed, usually 3, 4b (2-5); 6-C cephalad

of 7-C; 7-9-C usually subequal in length, 7-C 2,3b, 8,9-C normally single.

Trumpet: brown except for lighter extremes; distal portion gradually widened

from base; pinna medium-sized. Metanotum: lightly to moderately pigmented,

with haltere cases lighter; integumentary sculpturing absent; never more than

3 pairs of hairs developed. Abdomen: lightly to moderately pigmented, with

posterior portion of anterior segments and posterior segments lighter; seg-

ments II-VIII with weakly to moderately developed reticulate to imbricate in-

tegumentary sculpturing; smaller hairs concolorous with integument, larger

hairs darker; hair 5-I apparently single; 6,7-I moderately developed, usually

single (single, double); 1-II not dendritic, moderately developed, 3-5b; 2-II

cephalad of 3-II; 5-IV,V extending at least 0.75 distance to alveolus of hair 5

of following segment; 10-III-VII generally extending 0.5-0.7 distance to apex of

following segment; 10-III single or double; 10-IV usually 2, 3b (1-3); 1-VI usu-

ally mesad of hair 2, short and 4-7b; 1-VII mesad or laterad of hair 2; 9-VII

moderately developed, usually 6-8b (5-8), with longest branch extending 0. 6-

0.8 distance to alveolus of 9-VIII; 9-VIII strongly developed, usually 9-11b,

with longest branch extending 0.6-0.8 distance to apex of paddle. Terminal

70 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

Segments: male genital lobe projecting considerably beyond median caudal lobe.

Paddle: lightly pigmented, straw-colored, with midrib darker; longer than broad,

more or less oval in shape, distal portion of outer margin nearly straight; ex-

ternal buttress short, convex laterally, with minute spicules; midrib slight-

ly convex mesally; entire surface of paddle with small spicules.

LARVA (fig.15). Head: 0.95 mm. Siphon: 0.92 mm. Anal Saddle: 0.41

mm. Head: slightly broader than long; integument without ornamentation; lab-

rum short; largely light tan colored, lighter around imaginal eye, collar dark-

er; mental plate tan or brown, with 8-11 teeth on each side; hair 2-C apparent-

ly absent; 8-10-C usually 2, 3b (1-4), fine. Antenna: short; integument smooth;

uniformly light tan colored. Thorax: epidermal pigment strongly developed,

red in color; spicules absent; hairs strongly pigmented; hair 0-P 14-24b; 4-P

8-12b; 1-M moderately developed, 3-6b; 4-M double; 13-M 9-21b; 1-T moder-

ately developed, usually single (1-3b); 3-T 5-8b. Abdomen: segments VI and

VII with large sclerotized plates; hairs strongly pigmented; hair 1-I weakly de-

veloped, 2-5b; 2-I, II laterad of hair 1 of corresponding segment; 6-I, IJ multi-

ple; 10,11-I usually double (2, 3b); 13-I shorter than 10-I; 1-II moderately de-

veloped, 1-3b; 3-II,IV usually double (1-3b); 7-II 5-8b; 10-II 1-5b; 13-II 7-

14b; 1-III moderately developed, usually single (1-4b); 6-III-VI single; 7-III-V

7-15b; 1-VI weakly developed, 4-8b; 2-VI laterad of hair 1; 4-VI 2-4b; 7-VI,

VII usually 2, 3b (1-4); 13-VI 15-23b; 2-VII usually mesad of hair 1; 6, 8-VII 7-

12b. Segment VIII: sclerotized plate large, sometimes ringing segment; comb

scales normally in 1 row, 18-31 in number; individual comb scales fringed on-

ly, without a long apical spine; hair 3-VIII strongly developed, 9-12b. Siphon:

integumentary sculpturing moderately developed basally, imbricate; index about

3.9-4.2; extreme base dark brown, basal 0.6 light brown, fading to light tan in

apical 0.4; hair 1-S located about 0.53-0.61 from base of siphon, 10-13b,

strongly developed and with longest branches extending to apex of siphon. Anal

Segment: saddle complete; sclerotized incomplete ring present basad of saddle;

integumentary sSculpturing weakly developed, imbricate; saddle usually dark

brown at extreme base and sometimes at apex, remainder uniformly light brown;

hair 1-X usually 2, 3b (1-5), fine; ventral brush with 7 pairs of hairs; more

strongly developed hairs in ventral brush with 11-16 branches; gills mod-

erately long, dorsal pair 1.2-2.1 as long as ventral pair and 1.1-1.9 length of

anal saddle.

BIONOMICS. - According to Davis (1944:230), larvae are usually found in

living or dead bamboo internodes with either small or large openings and are

rarely found in bromeliad leaf axils. There is no information available on hab-

its of the adults.

SYSTEMATICS. This species is very distinct from all other Orthopodomy-

Adults are told from those of the Signifera group by having 4 narrow longi-

fda white lines on the mesonotum, broad flat decumbent scales on the ver-

tex, hindtarsal segment 1 longer than the hindtibia and no pra bristles. The

ee gcenitalia differ from the Signifera group in aedeagus shape and dentition,

by having 2 simple spiniforms and by having specialized mesal setae instead of

specialized tergomesal setae on the sidepiece. The larva of this species is the

only Orthopodomyia normally having a single row of comb scales.

The description of the adult of this species is based on specimens from the

state of Sao Paulo, Brazil; specimens from Venezuela agree in almost all de-

tails but differ by sometimes having the mesonotal integument more evenly red-

dish-brown, coxae and trochanters usually without tannish or brownish scales,

hindtarsal segment I sometimes white speckled and the fan of scales on tergite

1 sometimes all white. In addition, the markings on the proboscis are appar-

Zavortink: Genus Orthopodomyia 71

ently somewhat different in the Venezuelan material, the female having a long

continuous or broken dorsal streak of white scales while the male lacks even a

short streak in the same position. The descriptions of the larva and pupa are

based on material from Venezuela. I have seen 1 larva, possibly from Brazil,

which appears to agree with the Venezuelan specimens (it is in very poor con-

dition) except for having 13 or 16 teeth, instead of 8-10, on each side of the

mental plate.

This is apparently a relict species restricted to a few localities in South

America. Specimens have been collected only near Caracas, Venezuela and

Rio de Janeiro and Sao Paulo, Brazil. It is evidently at least distantly related

to the Signifera group since the adult ornamentation, especially on the thorax,

is Similar and apomorph in both groups. However, the male genitalia are not

at all similar to the Signifera group; the aedeagus is remotely similar to the

Madagascan Vernoni group, whereas similarly developed specialized setae on

the mesal surface of the sidepiece are found only in albipes and madrensis,

members of the Oriental Albipes group. Two spiniforms, the rule in albicosta,

are found as anomalies in both the Vernoni and Albipes groups. The pupa is

similar to both the Signifera group and the Thomasina section; the development

of hairs 3-C, 10-IV and 1-VI and the presence of spicules over the entire pad-

dle surface are notable similarities to fascipes, while hairs 10-12-C and 1-II,

III are like those of the Signifera group. The larva is similar to the Signifera

group in head shape, in lacking ornamentation on the antenna, in development

of hairs 8-P, 1-I-V, VII, and 13-I, in position of hair 2-I, II-VI and in siphon

shape and ornamentation, but is similar to phyllozoa in the development of hairs

1,4-M and 3-T.

DISTRIBUTION. Discontinuous, found only in north-central Venezuela

and southeastern Brazil. Material Examined: 75 specimens; 260, 2192, 13P,

15 L; 13 individual rearings (7 larval, 5 pupal, 1 incomplete).

BRAZIL. Guanabara: Rio de Janeiro, Muda da Tijuca, 29 June 1953, H.

Ferreira, 1 ¢ [IOC].

Sao Paulo: Boraceia, Feb 1961-Aug 1962, O. P. Forattini, 30°, 49 [FH].

Cantareira, 16 Apr 1905, 12 [BMNH]. Cochocirinha, 22 Nov 1904, 1% [BMNH].

No locality or date: 2 9 [BMNH].

VENEZUELA. Aragua: Maracay, 1 Dec 1967, R.I.C. Hansell and J.J.

Vera (VZ 83), 3 lpo (83-10,14, 70), 4 lp? (83-11,12,15,16), 2 pc (83-100,

103), 3 p? (83-17, 107, 108), 1 1p (83-20), 4 L [UCLA]. Ocumare de la Costa, ©

14 July-18 Aug 1927, M. Nunez Tovar, 140, 59, 1 L [USNM], 1° [BMNH].

Villagos, 9 Aug 1927, M. Nunez Tovar, 1 % [USNM].

No locality: March 1939, P.J. Anduze (51), 1 L [USNM].

NO DATA. 1, 12 [USNM]; 1 L [BMNH]. 7

Additional records from the literature. BRAZIL. Rio de Janeiro: Tereso-

polis, May 1942-Sept 1943, L (Davis, 1944:230).

THOMASINA SECTION

FEMALES. Head: integument of head capsule, clypeus, torus and flagel-

lum brown to dark brown; eyes Separated above antennae by a row of white to

cream-colored decumbent scales; frontal bristles amber to dark brown; orbital

bristles amber to dark brown, usually 2 lateral pairs and 2 or 3 mesal pairs;

ocular border absent; vertex and occiput with very numerous narrow curved

decumbent scales and very numerous erect scales, decumbent scales broader

and denser in front of dorsolateral scales and mesally; dorsolateral scales

72 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

brown anteriorly; suborbital bristles pale; clypeus an elongate angular lobe,

bare; labium slightly swollen distally, shaft bristles absent, largely brown- or

black-scaled with white scales forming a small to moderate-sized dorsal patch

0.7-0.8 distance from the base and a small dorsoapical patch, and with numer-

ous scattered yellow and sometimes white scales; palpus straight, 0.36-0.38

of proboscis, 4-segmented, segment 4 short, segments 2 and 3 largely brown-

or black-scaled with numerous scattered yellow and sometimes white scales or

largely mixed dark- and light-scaled, apex of segment 3 and segment 4 with

silvery-white scales; inner half of torus with small scales except for bare spot

mesally and dorsal portion of inner half with short, fine hairs; flagellar seg-

ment 1 not noticeably swollen and with a few small scales; dorsal flagellar bris-

tles about 1.5-2.0 length of their segment on proximal segments, ventral bris-

tles absent on segments 1-6 or 7. Thorax: paratergite narrow to moderately

broad, bare; posterior dorsocentral bristles usually not in a double row; hu-

meral, lateral prescutal and posterior fossal bristles not numerous enough to

completely enclose fossa; fossal bristles absent; mesonotal bristles dark

brown, parascutellar and scutellar bristles lighter; mesonotal ornamentation

consisting of light and dark scales of various sizes and densities almost com-

pletely covering the scutum, as follows: (1) acrostichal scales in a broad band

accompanying the acrostichal bristles, not separated by a narrow bare longi-

tudinal area from the dorsocentral scales, (2) anterior prescutellar scales in

a more or less crescent-shaped or triangular area anterior to prescutellar

space, small and large, yellow or golden or with a few brown posteriorly, (3)

dorsocentral scales in a broad band accompanying the dorsocentral bristles,

(4) lateral prescutellar scales accompanying the lateral prescutellar bristles

and extending caudad to margin of scutellum, large, brown anteriorly, (5) lat-

eral prescutal scales extending from anterior promontory to scutal angle along

edge of mesonotum, small, (6) posterior fossal scales extending from scutal

angle to dorsocentral scales anterior of scutal suture, small, usually silvery-

white and yellow or golden, sometimes brownish, (7) fossal scales in the area

enclosed by the dorsocentral, lateral prescutal and posterior fossal scales, -

small, (8) supraalar scales extending from scutal suture to parascutellum

among and mesad of supraalar bristles and with middle extension ventrad to

margin of paratergite, scales mostly small mesally and large laterally; para-

scutellum usually with patch of white to silvery-white scales; lateral and mid-

scutellar lobes each with an extensive patch of large scales which extend over

the caudal margin of the scutellum; larger apn, ppn and pra bristles brown,

remaining pleural bristles lighter; upper anterior apn bristles 2-5 strongly de-

veloped, 2 or 3 weakly to moderately developed, lower apn bristles usually ab-

sent; ppl bristles 3-5 strongly developed, 3-6 weakly to moderately developed;

pra usually with 1 bristle (0-2); upper stp bristles 1, strongly developed, pos-

terior stp bristles 4-11, moderately to strongly developed, lower stp bristles

2-6, weakly to moderately developed; upper posterior mep bristles 4-10, mod-

-erately developed, anterior mep bristles absent; pleural scaling consisting of

extensive areas and patches of scales; apn with narrow decumbent scales in an

upper anterior patch and narrow and broader decumbent scales in a ventral

diagonal row; ppn scaled in posterior-ventral half, scales above narrow and

curved, those below broad and flat; psp scales absent; pra with scales on an-

terior edge and below knob; ppl with broad flat and semierect scales in a dense

patch around the bristles; entire pst with a dense covering of broad flat scales;

pex, ssp and hypostigial scales present, broad; upper half of stp completely

covered with broad flat scales except for a small bare area below the ssp

scales and around the upper stp bristle, lower half with a posterior ventral row

Zavortink: Genus Orthopodomyia 73

of broad flat and semierect scales; mep with elongate semierect scales anterior

to the upper posterior bristles and with broad flat scales in an upper anterior

patch continuous with the patch of semierect scales; a few additional large

brownish or transparent scales may be found scattered on stp, mep and the

meron. Legs: integument of coxae same color as pleuron or upper portion,

especially on forecoxa, lighter; forecoxal bristles amber, midcoxal bristles

usually brown and hindcoxal bristles usually brownish-amber; forecoxa with

broad flat scales on anterior surface and among and behind bristles and in a

small patch on lower posterior surface, midcoxa with broad flat scales among

and anterior to bristles, hindcoxa with broad flat scales on anterior surface;

trochanters with brown, white and sometimes yellowish flat scales on anterior,

inner and posterior surfaces; femora with small white or yellowish knee spots;

hindtarsal segment 1 longer than hindtibia; hindclaws slightly smaller than

fore- and midclaws. Wing: remigial bristles absent; vein Rg 1.85-2.17 length

of vein R9+3; vein My49 1.58-2.11 length of vein M distad of crossvein m-cu;

plical area with light scales; alula scales moderately broad; upper calypter

with bristles; scales symmetrical and asymmetrical, moderately to very broad,

mostly spreading, some large and cordate; wing largely brown- or black-scaled,

with light scales forming small patches and scattered, as follows: (1) costal

vein with small basal and humeral light patches, sometimes with small subcos-

tal light patch and with few to numerous scattered light scales, (2) vein Sc with

few to numerous scattered light scales and sometimes with small sectoral and

subcostal light patches, (3) vein R with normally small basal, humeral and sec-

toral light patches and sometimes with scattered light scales, (4) vein Ry with

accessory sectoral light patch and few to numerous scattered light scales, (5)

veins Rg, R445, Cu and 1A usually with basal light patch and vein M sometimes

with basal light patch, (6) vein R943 with light patch opposite base of vein R45,

(7) vein Cu usually with light patch over furcation, (8) veins M and Cu usually

with light patches distad of crossveins, (9) veins Rg and Rg usually with an api-

cal light patch, (10) veins Rg, M, Cu and 1A with few to numerous scattered

scales. Abdomen: integument of tergites and distal sternites light to dark

brown, proximal sternites evenly light to dark brown or with much lighter, even

white, areas beneath light scales; laterotergite with numerous silvery-white

scales; tergite 1 with entirely or largely dark brown- or black-scaled fan and |

with numerous scattered light and dark scales; remaining segments with dark

scales brown or black, light scales largely white, but some yellowish; tergites

largely dark-scaled, tergites II-VII with long straight or curved diagonal light

patch starting basolaterally, tergite VIII with basolateral light patch; some or

most tergites with few to numerous yellowish scales basally and/or apically;

sternite VIII bare mesally, dark-scaled laterally. |

FEMALE GENITALIA. Tergite IX with 1 pair of dorsolateral setae.

MALES. Similar to females. Labium: strongly sclerotized and much

greater in diameter throughout length than in female; upper portion bare of

scales, ventral portion largely dark brown- to black-scaled, white scales ina

small dorsoapical patch, yellow or golden scales scattered. Palpus: 0.90-0.93

of proboscis, segment 5 moderately long, segment 3 with moderately developed

apical bristles; largely yellowish- and white-scaled with a few brown scales or

largely mixed brown- and light-scaled, white or silvery-white scales usually

near base of segment 1 and at joints between segments 2-3, 3-4 and 4-5, seg-

ment 4 often with fewer light scales than segments 2 and 3, segment 5 silvery-

white- and brown-scaled or entirely silvery-white-scaled. Antenna: integument

of flagellum brown to dark brown except for lighter areas basad of bristles on

shorter segments; segment 13 about 1.5 length of segment 12; torus not as en-

74 | Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

larged as in males of other groups; flagellar segment 1 with small tuft of mod-

erate-sized scales; flagellar bristles not as strongly developed as in males of

other groups, bristles about 0.2-0.3 length of entire flagellum on proximal seg-

ments and 0.3-0.5 length of segment 13 on that segment. Legs: anterior fore-

and midclaws with basal tooth large, visible in pinner specimens, posterior

fore- and midclaws simple. Abdomen: tergites often with narrow basal and

apical yellow or golden bands; sternites with more extensive light-scaling;

sternite VIII dark-scaled vith basolateral ges patch; sidepiece with light and

dark scales.

MALE GENITALIA. Segment VIII: posterior margin of tergite without lobe,

straight or slightly rounded mesally. Segment IX: tergite somewhat reduced,

without fine setae dorsolaterally. Sidepiece: shape varied, basal mesal lobe

with 2 or 3 very strong, short, blunt and flattened (at least distally) apical bris-

tles tergally and with 1-8 long fine bristles sternally; mesal surface membra-

nous for a considerable distance distad of basal mesal lobe; mesal and tergo-

mesal surfaces with or without specialized bristles. Spiniform: 1, with apex

flattened, broadened, partially rolled and divided into numerous more or less

finger- -like lobes. Aedeagus: weakly to moderately sclerotized, varied in form.

Proctiger: basolateral sclerotization very strongly developed; paraproct scle-

rotization with 2-4 apical teeth; cercal setae 2-5.

PUPAE. Known only for fascipes.

LARVAE. Known completely only for fascipes.

BIONOMICS. The immature stages are found in treeholes, bamboo inter-

nodes and artificial containers. Females are not known to bite man.

SYSTEMATICS. This section contains the 2 species fascipes and sampaioi.

It is well marked in the adult stage and the male genitalia. The immature ©

stages are adequately known only for fascipes.

The affinities of the Thomasina section are no more obvious than those of

the other sections and groups of Orthopodomyia. Edwards (1932:108) placed

fascipes in his Group A (Orthopodomyia) on the basis of adult ornamentation.

However, characteristics of the male genitalia, larva and pupa indicate that

fascipes is not closely related to the other species Edwards placed in that group.

The male genitalia of the Thomasina section have several unique features, but

the spiniform is like that of the Holarctic Signifera and the Madagascan Vernoni

groups. The pupa of fascipes resembles the Signifera group while the larva

shares characteristics with both the Signifera group and the Central American

Folicolae section. Since it is possible that some of the larval and pupal features

of fascipes have been acquired by hybridization, caution should be exercised in

judging the affinities of the Thomasina section on characteristics of the imma-

ture stages until those stages are known for sampaioi.

Males of fascipes and sampaioi have fewer and shorter flagellar bristles

and smaller tori than males of other Orthopodomyia. These differences indi-

cate that the auditory function of the antennae is less well developed in these

species and imply a difference in mating behavior. The proboscis of the males

of these species is also unusual in structure. It is very stout, nearly equal to

the forefemur in diameter, and strongly sclerotized. The function of this spe-

cialized organ is unknown.

Although fascipes and sampaioi are basically allopatric, their ranges ap-

parently meet or overlap in the Brazilian state of Goias. Larvae which I have

seen from the highlands separating the Parana and Tocantins drainage systems

are indistinguishable from f ascipes, yet the few adults available from the same

area are sampaioi. It is possible that the 2 species meet along the entire arc

of highlands which separate the Parana River system from the nee and east-

ward draining rivers of South America.

Zavortink: Genus Orthopodomyia fee

DISTRIBUTION. Central and South America.

KEYS TO SPECIES

Adults

1. Femora, tibiae and tarsal segments 1 with distinct light patches or irreg-

ular rings; acrostichal scale band usually with large cream-colored or

yellowish scales at anterior promontory and in tufts throughout length of

SCAle DANO. cas) Mueaei Ue bags a we ak eee es ee Les

Femora, tibiae and tarsal segments 1 without distinct light patches or ir-

regular rings; acrostichal scale band with large white or cream-colored

scales in more or less complete row from anterior promontory to caudal

end Of SCAG DANG oo. ee ee lea WH eee a Ee ey oe Be eo

Male Genitalia |

1. Sidepiece long conical, not strongly constricted distally; basal mesal lobe

with only 1 or 2 long fine bristles sternad of strong blunt bristles; aede-

agus with 1 long, strong process on each half sternally. . . 8. fascipes

Sidepiece broad basally, strongly constricted distally; basal mesal lobe

with 6-8 long fine bristles sternad of strong blunt bristles; aedeagus

without strong sternal processes, but with a sclerotized tooth along

mesal édge of each half sternaily so. is eos bee ee eee Oy Samapaiol

8. Orthopodomyia fascipes (Coquillett)

Figs. 16,17

1905. Mansonia fascipes Coquillett, 1905:182-183. *TYPE: lectotype ¢ (338b),

pathway to Las Loras, Rio Aranjuez, near Puntarenas, [Puntarenas],

Costa Rica, adult found in the crevice of a tree trunk, 13 Sept 1905, F.

Knab; selection of Stone and Knight (1957:198) [USNM, 8296].

1910. Mansonia longipalpis Newstead and Thomas, 1910:145-147. *TYPE: lec-

totype , specimen labeled ''Type ¢,'' Bach's outside house, Manaos,

Amazonas, Brazil, adult captured from 4 to 6 PM, 23 Aug 1906; PRES-

ENT SELECTION [BMNH]. Synonymy with fascipes by Howard, Dyar and

Knab (1917:882-883). : | |

1935. Orthopodomyia (Orthopodomyia) townsendi Lima, 1935:178. *TYPE: ho-

lotype 3, Rio Tapajos, Boa Vista, Para, Brazil, 11 Oct 1932, C.H.T.

Townsend [IOC, vial 582]. Synonymy with fascipes by Lane (1951:335).

1958. Orthopodomyia bacigalupoi Martinez and Prosen, 1958:17-22. TYPE: ho-

lotype &, Buenavista (Tacu), Ichilo, Santa Cruz, Bolivia, reared from a

larva found in a treehole, Apr 1955, A. Martinez and A. F. Prosen

[MEPRA]. NEW SYNONYMY.

Orthopodomyia fascipes of Howard, Dyar and Knab (1917:882-886); Bonne-Wep-

ster and Bonne (1923:127); Dyar (1923a:183); Dyar and Shannon (1924:484);

Bonne and Bonne-Wepster (1925:490-494); Dyar (1925:179; 1928:396-397);

Lima (1930:255); Shannon (1931:9); Edwards (1932:108; 1934:632); Town-

send (1934:497); Komp (1936:66); Antunes (1937:79); Chagas, Castro and

Ferreira (1937:385); Kumm and Novis (1938:511); Lane (1939:96-97); Sene-

vet and Abonnenc (1939:588-590); Kumm, Komp and Ruiz (1940:405, 417, 418);

76 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

Anduze (1941:12); Floch and Abonnenc (1942:5); Senevet (1946:326); Anduze

(1947:359-360); Floch and Abonnenc (1947:7); Roth (1948:168-169); Arnett

1949:235-236); Bates (1949:23); Arnett (1950:107, 109,111); Bohart (1950:

403, 404); Knight and Mattingly (1950:2); Galindo, Carpenter and Trapido

(1951:127); Lane (1951:335); Carpenter and Peyton (1952:676); Levi-Castillo

(1952:555); Lane (1953:620-622); Levi-Castillo (1953:37); Galindo, Carpen-

ter and Trapido (1955:159, 161); Trapido, Galindo and Carpenter (1955:535);

Stone, Knight and Starcke (1959:123); Aitken (1960:4); Cerqueira (1961:143);

Fauran (1961:25); Maciel (1962:489); Cova-Garcia, Sutil and Rausseo (1966a:

53; 1966b:51-52, 117, 143, 371).

Orthopodomyia (Orthopodomyia) fascipes of Lima (1935:176, 177, 178).

Bancroftia fascipes of Dyar and Knab (1910:264).

Mansonia fascipes of Coquillett (1906:25); Dyar and Knab (1906:185); Knab (1907:

153); Aiken (1909:13-14); Theobald (1910b:447, 451); Stone and Knight (1957:

198).

Orthopodomyia bacigalupoi of Stone, Knight aa Starcke satis 123); Prosen,

Carcavallo and Martinez (1964:90).

Orthopodomyia townsendi of Lane (1939:97); Knight and Mattingly (1950:2).

Orthopodomyia aff. fascipes of Townsend (1934:497).

Thomasina longipalpis of Newstead and Carter (1911:553-556).

FEMALE. Wing: 4.00 mm. Proboscis: 2.52 mm. Forefemur: 2.37 mm.

Abdomen: about 2.7 mm. Head: decumbent scales yellow or yellow and white

mesally, yellow, white and brown laterally, erect scales usually yellow mesal-

ly, largely brown laterally; posterior dorsolateral, lateral and ventral scales

white or yellow-tinged; palpus 0.38 of proboscis; torus and flagellar scales

dingy. Thorax: integument of mesonotum reddish brown to dark brown lateral-

ly, anterior mesal portion usually lighter and posterior mesal portion usually

darker, scutellum lighter than lateral portion of mesonotum, postnotum darker

except for lighter lateral edge, pleuron reddish-brown to dark brown dorsoan-

teriorly, usually lighter ventrally and posteriorly; acrostichal scale band with

large cream-colored or yellowish scales at anterior promontory and in tufts

throughout length of scale band or rarely in a more or less complete row

throughout length of band, brown scales at caudal end, remaining scales large

and small, yellow or golden; dorsocentral scale band with small white and gold-

en or brown scales at anterior promontory, large cream-colored or rarely

golden scales in a tuft at caudal end or rarely in a row throughout the posterior

portion of the scale band, remaining scales large and small, mostly yellow or

golden; lateral prescutellar scales silvery-white to yellowish posteriorly; lat-

eral prescutal scales silvery-white anteriorly and sometimes posteriorly, usu-

ally yellow or golden posteriorly; fossal scales usually mostly yellow or gold-

en, sometimes brownish; supraalar scales silvery-white anteriorly and in ven-

tral extension, silvery-white or silvery-white and brown among bristles, re- |

mainder yellowish; scutellar scales white to silvery-white or yellowish; pleural

scales creamy-white to yellow-tinged or dingy. Legs: forecoxal scales white

or yellow-tinged, mid- and hindcoxal scales white or yellowish above, brown

or brown and yellowish below; forefemur largely brown- or black-scaled ex-

cept for white and/or yellow scales in a small patch distad of every bristle, in

an irregular more or less complete ring about 0.7-0.8 distance from base, and

sometimes scattered; midfemur like forefemur except that base of posterior

surface has short to long pale streak; hindfemur entirely white- or yellowish-

scaled in ventral portion of anterior surface and basal 0.4-0.6 of posterior

surface and largely white- or yellowish-scaled in distal portion of anterior

Zavortink: Genus Orthopodomyia 17

surface, remainder brown-scaled with white and/or yellowish scales in small

patches and scattered; tibiae largely brown- to black-scaled, with white, dingy

or yellowish scales in numerous irregular patches and rings, at apex, scat-

tered, in streak on ventral and anterior surface of hindtibia and sometimes in

streak on posterior surface of foretibia; tarsi largely brown- or black-scaled,

segment 1 with white to yellowish scales forming small dorsobasal patch or

narrow ring on foretarsus, narrow basal ring on midtarsus, small dorsobasal

patch on hindtarsus and irregular patches or speckles on midtarsus and some-

times fore- and hindtarsi; foretarsal segments 1-3 or 4 usually with a few

white scales at apex, segment 2 sometimes with small light patches, segment

3 sometimes white- or cream-scaled at base or over entire dorsal surface;

midtarsus usually with white to cream-colored scales covering apex of segment

1, ends of segment 2, ends or entire surface of segment 3, entire dorsal sur-

face of segments 4 and 5, sometimes in speckles or small patches on segment

2 and sometimes streaked on ventral surface of segment 1; hindtarsal segment

1 with or without a few light scales at apex, segment 2 with white scales form-

ing narrow to very broad apical ring and sometimes small patches or speckles,

segments 3-5 entirely white-scaled or more commonly segment 3 with dark-

scaled basal and/or preapical ventral patch or complete ring, segment 4 dark-

scaled except at ends and segment 5 dark-scaled ventrally. Wing: light scales

in small patches usually white, sometimes dingy yellow, scattered light scales

mostly dingy yellow, some white; veins Ro,9 and M usually with distinct light

patch over furcation; vein R4.5 rarely with apical light patch. Haltere: integ-

ument beige to brown; scales transparent to whitish and/or brownish. Abdo-

men: sternites II and III with large basolateral light-scaled patches, mesal and

apical portions of segments dark-scaled, sternites IV-VII largely dark-scaled,

with light scales in basolateral patch or weakly developed basal band.

MALE. Similar to female. Palpus: 0.90 of proboscis. Antenna: flagellar

— scales dingy.

MALE GENITALIA (fig. 16). Segment VIII: posterior margin of tergite

straight or slightly rounded mesally. Sidepiece: cylindrical to long conical;

basal mesal lobe with 1 or 2 long fine bristles separated by a gap from more

tergal strong blunt bristles; mesal surface without specialized bristles; tergo-

mesal surface usually with 1-7 short, strong, flat and curved bristles distad of

basal mesal lobe. Aedeagus: more or less parallel sided; sternal surface with

2 long, strong processes; base with irregular projection between or below ven-

tral parameres. Proctiger: paraproct sclerotization with 2-4 apical teeth;

cercal setae 2-4.

PUPA (fig.16). Abdomen: 4.38 mm. Trumpet: 0.70 mm. Paddle: 0.93

mm. Cephalothorax: moderately pigmented, straw-colored to tan, with upper

and posterior portions of wing case, portion of mesonotum caudad of trumpet,

and sometimes middorsal ridge darker; integumentary sculpturing strong, re-

ticulate; hairs largely concolorous with integument; all hairs shorter than

trumpet; hairs 1,7-C moderately developed, usually subequal, usually double

(1-3b); 2-C short, single or double; 3-C very weakly developed, usually 3, 4b

(2-5); 4-C slightly shorter than 1-C, usually double (1-3b); 5-C short, usually

3-5b (2-6); 6-C cephalad of 7-C; 8,9-C short, usually single or 8-C 2, 3b.

Trumpet: brown except for extreme base and apex; distal portion gradually

widened from base; pinna large. Metanotum: uniformly moderately pigmented,

tan; integumentary sculpturing strong, reticulate; never more than 3 pairs of

developed; hair 10-C moderately developed, usually 3-5b (2-6). Abdomen: an-

terior segments moderately pigmented, tan, posterior segments lightly pig-

mented, straw-colored; anterior segments darker cephalad and along posterior

78 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968 ©

edge; anterior segments with strongly developed imbricate integumentary sculp-

turing, posterior segments with weaker sculpturing; smaller hairs concolorous

with integument, larger hairs darker; hair 1-II usually not dendritic, usually

10-13b; 2-II cephalad of 3-II; 5-IV, V reaching to or nearly to alveolus of hair

5 of following segment and normally single; 10-III-VII short, not extending be-

yond 0.5 distance to apex of following segment; 10-III, IV usually 2, 3b (1-4);

1-VI, VII mesad of hair 2 of corresponding segment; 1-VI short, usually 6, 7b

(5-10); 9-VII moderately developed, usually 4-6b, longest branch about 0. 3-

0.5 distance between alveoli of hairs 9-VII and 9-VIII; 9-VIII strongly devel-

oped, usually 7-9b, longest branch not extending beyond end of external but-

tress of paddle. Terminal Segments: male genital lobe projecting considerably

beyond median caudal lobe. Paddle: lightly pigmented, straw-colored, with

midrib darker; longer than broad, usually somewhat angular apically, with dis-

tal portions of inner and outer margins straight; external buttress long, slight-

ly convex laterally, with minute spicules; midrib slightly convex mesally; en-

tire surface of paddle covered with small spicules.

LARVA (fig.17). Head: 1.13 mm. Siphon: 1.38 mm. Anal Saddle: 0.43

mm. Head: slightly broader than long; integument spiculose; largely straw-

colored, lighter around imaginal eye, darker posteriorly, collar brown; labrum

short; mental plate tan, usually with 10-12 teeth on each side; hair 2-C absent.

Antenna: short; integument with small spicules; uniformly straw-colored; hair

1-A moderately developed, 6-8b (4-9). Thorax: epidermal pigment moderately

to strongly developed, red to purple in color; spicules not conspicuous; hairs

strongly pigmented; hair 9-P strongly developed, long, usually 7-9b; 1-M mod-

erately developed and usually 3-5b in non-hairy form, strongly developed and

up to 20b in hairy form; 4-M always single; 1-T long and single in non-hairy

form, long and up to 20b in hairy form; 13-T strongly developed, long, usually

5-10b. Abdomen: segment VI with or without dorsal sclerotized plate, segment

VII usually with large, often quadrangular, sclerotized plate; hairs strongly

pigmented; hair 1-I usually 3-5b and weakly developed in non-hairy form, up

to 15b and strongly developed in hairy form; 2-I, II laterad of hair 1 of corres-

ponding segment; 6-I, II multiple, 6-II-VI single or up to ob in hairy form;

13-I shorter than 10-I; 1-III-V long, single or double in non-hairy form, up to

18b in hairy form; 13-III, IV long, usually single or double in non-hairy form,

up to 18b in hairy form; 5-IV,V single and longer than hair 3 of corresponding ~

segment in non-hairy form, up to 10b and longer than or subequal to hair 3 in

hairy form; 13-V long, usually 2-4b in non-hairy form, up to 11b in hairy form;

1-VI weakly developed in non-hairy form, strongly developed in hairy form;

2-VI, VII laterad of hair 1 of corresponding segment; 5-VI longer than 3-VI ex-

cept in some hairy individuals. Segment VIII: large sclerotized plate present;

comb scales in 2 rows; anterior comb scales usually 19-26, posterior comb

scales usually 9-14; larger anterior comb scales shaped like posterior comb

scales, with 1 long apical spine and inconspicuous fringe in basal portion. Si-

phon: index about 3.5-6.0; uniformly light brown to dark brown except for dark-

er base and lighter apex or with portion distad of alveolus of hair 1-S somewhat

darker; integumentary sculpturing strongly developed, imbricate; hair 1-S lo-

cated about 0.3-0.4 from base of siphon, longest branches extending about 0. 6-

0.9 distance from alveolus to apex of siphon. Anal Segment: saddle complete,

largely light brown to dark brown, base and dorsal surface usually darker; in-

tegumentary sculpturing strongly developed, imbricate; sclerotized band or

incomplete ring present at base of saddle; hair 2-X usually with 20-28b; ven-.

tral brush usually with 8 pairs of hairs, the more strongly developed hairs with

about 16-23 branches; gills moderately long, dorsal pair about 1.2-1.6 as long

Zavortink: Genus Orthopodomyia 719

as ventral pair and 1.0-2.6 length of anal saddle.

BIONOMICS. The immature stages have been collected in treeholes, bam-

boo stumps, open and closed-top bamboo traps, cement containers and cess-

pools. Adults have been captured in light traps, flying, and resting on tree-

trunks and buildings. :

SYSTEMATICS. Fascipes is the dominant, widespread species of Ortho-

podomyia in tropical America. It has 3 synonyms, longipalpis, townsendi, and

bacigalupoi. Newstead and Thomas were apparently unacquainted with Coquil-

lett's 1905 description of fascipes when they described longipalpis in 1910.

Howard, Dyar and Knab (1917:882) first synonymized longipalpis with fascipes

and their treatment has been accepted by all subsequent authors.

Townsend (1934:497) reported collecting a large series of fascipes and 1

male and 1 female of ''Orthopodomyia aff. fascipes (?)'"' at Boa Vista, Para,

Brazil. The latter differed from fascipes by the following features: (1) the

structure of the male genitalia, (2) tergites II-VI of the female pale yellowish,

and (3) the fourth hindtarsal segment only slightly longer than the fifth and en-

tirely white-scaled in both sexes. The male was sent to the United Stated Na- |

tional Museum and identified as fascipes by Dr. Alan Stone. I have examined

the specimen (labeled "Orthopodomyia n. sp." by Townsend) and agree with

Stone. The differences between the male genitalia of fascipes and ''Orthopodo-

myia aff. fascipes (?)'' observed by Townsend are artifacts caused by flatten-

ing of the slide mounted preparations. The female of ''Orthopodomyia aff. fas-

cipes (?)' was sent to Cesar Pinto. It was examined, considered to be distinct

from fascipes on the basis of the hindtarsus, and described as townsendi by Li-

ma in 1935. Lane (1951:335) reduced townsendi to synonymy with fascipes and

I concur with that interpretation. The hindtarsus of the holotype of townsendi

is not unique in ornamentation or structure. Measurement of segments 4 and

0 has shown that their relative lengths are no different thanin fascipes. As

pointed out by Howard, Dyar and Knab (1917:886), the tarsal ornamentation of

fascipes is quite variable and the last 3 segments may be entirely white-scaled

on all legs. Since all tergites of the holotype are denuded of scales, it is im-

possible to verify whether II-VI were pale yellowish as stated by Townsend. |

Martinez and Prosen described bacigalupoi in 1958 from material collected

in Bolivia. Since the original description of the species does not compare baci-

galupoi with any other species of Orthopodomyia, it is not clear why these au-

thors considered their material to represent an undescribed species. I have

examined 2 male paratypes and have found that they agree with fascipes in or-

namentation and male genitalia and as a paneeanenes. Iam synonymizing baci-

galupoi with fascipes. :

Adults of fascipes are easily distinguished from Ssampaioi by the following:

(1) distinct light patches or irregular rings on all femora, tibiae and first tar-

sal segments, (2) fewer (usually, see below) large outstanding white or cream-

colored scales in the acrostichal scale band and the posterior portion of the

dorsocentral scale band, and (3) yellower or dingier color of many of the light

scales on all parts of the body. In addition to the characters given in the key,

the male genitalia of fascipes and Sampaioi differ in shape of the aedeagus. The

larvae of fascipes and sampaioi differ by at least the features indicated in the

description of sampaioi. ,

The wing and tarsal ornamentation of fascipes are quite variable. While

some light patches of the wing are always present and distinct, others may be

present and distinct, present but obscured by scattered light scales, or absent.

The scattered light scales vary in number and color. As indicated above, the

last 3 tarsal segments may be entirely white-scaled on all legs, but more com-

80 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

monly at least some of the segments involved have dark scales (see description).

The acrostichal scale band of fascipes usually has tufts of large white or cream-

colored scales throughout its length, but in a few individuals there is an almost

complete row of these scales. In this respect these individuals resemble sam-

paioi. The same type of variation is found in the posterior portion of the dor-

socentral scale band; most individuals have only a single tuft of large light

scales directly in front of the brown lateral prescutellar scales, but a few have

a long row of large outstanding light scales.

Both non-hairy and hairy forms of the larva are known. Extremely hairy

individuals may have hairs which are single in the non-hairy form with as many

as 20 branches, and even hairs 6-JII-VI may be 3-5 branched. All pupae, in-

cluding those from extremely hairy larvae, are non-hairy.

Fascipes is sympatric with phyllozoa in the northern portion of its range

and some of its characteristics may have been obtained from that species. The |

spotted legs and yellower coloration of the adult and the multiple hair 1-M and

spiculate head capsule and antenna of the larva are characteristics of fascipes

found in phyllozoa but not in sampaioi.

Fascipes is apparently sympatric with albicosta in Venezuela and in con-

tact with sampaioi in Brazil. It may be sympatric with kummi in Central Amer-

ica. The pupa of fascipes is near that of both albicosta and kummi and possibly

this similarity is due to hybridization with one of these species.

DISTRIBUTION. Nicaragua, south to Bolivia and the Brazilian states of

Goias and Minas Gerais. Material Examined: 1163 specimens; 199 o, 271 9,

1 gynandromorph, 7 isolated “ genitalia, 169P, 516L; 118 individual rearings

(97 larval, 14 pupal, 7 incomplete). —

BOLIVIA. Santa Cruz: Ichilo, Buenavista, Tacu, A. Martinez (955), 2¢%

[MART ].

BRAZIL. Amazonas: Manaus, 23 Aug 1906, 2 lectotype of longipalpis

[BMNH]; 4 Nov 1910, H. W. Thomas, Po ([BMNH]. No locality or date, 2c,

2° [BMNH].

Bahia: Piraja, 8 Oct 1929, R.C. Shannon, 1 % genitalia, 2P, 7 L [USNM];

1 L[USNM]. Rio Coruripe, 13 Jan-June 1931, 20°, 29 [BMNH]. No locality,

June 1931, 3 [BMNH]. No locality or date, 1“ [BMNH].

| Goias: Bomfim, 1 L[USNM]. Inhumas, 2 L [USNM].

Para: Belem, i Apr 1941, W.H. W. Komp: i? [UCLA]. Curralinho, Rio

Pagdo (K1184), 4 o, 2e76 B [USNM], 3 L[BMNH]. Curralinho (K1436), 2,

1? [BMNH], 20 [USNM]. Boa Vista, Rio Tapajos, 11 Oct 1932, C.H.T. Town-

send, ? holotype of townsendi [IOC]; C.H.T. Townsend, 150, 8 2 [USNM].

Piaui: Teresina, July 1934-Mar 1935, 10%, 19 [CU]; 7 L [USNM].

-~COLOMBIA. Boyaca: Muzo, 18 July 1936, E. Osorno, 1 ¢ [CU].

Meta: Restrepo, July-Aug 1935, W. H. W. Komp, 1G, 39, 1° genitalia,

3 L[USNM]. Villavicencio, Retiro, Aug 1935 (KO3-6), 3 ot, 2 [UCLA].

Valle del Cauca: Buenaventura, 3 Sept 1964, V.H. Lee (COL 3), 3: 1lpo

(3-10.11, 14), Pip? (3-12), 1 Pp 3-17), 1 L (UCLAI.:

No locality or date: E. Osorno, 1% [UCLA].

COSTA RICA. Bananito, 1% [USNM]. Golfito, Punta Rieles, 17 Mar 1943,

T.H.G. Aitken, 19 [USNM]. Osa Peninsula, Rincon, 29 June 1963, C.L.

Hogue (CR 130), 1 lp (130-502), 1 lp? (130-501), 1 L [UCLA]; same data

(CR 131), 5 lpo (131-301-303, 306, 307), 2 lp? (131-304, 305), 4 L [UCLA].

Puntarenas, Rio Aranjuez, F. Knab (238), 2 lectotype of fascipes, 60°, 3 9

USNM].

ECUADOR. Napo: Coca, Coca and Napo Rivers, 23 Apr-12 May 1965,

L.E. Pefia (ECU 8), 19 [UCLA]. Pompeya Island (Napo River), 19-26 May

Zavortink: Genus Orthopodomyia |. 81

1965, L.E. Pefia (ECU 19), 19 [UCLA].

FRENCH GUIANA. Cayenne: aise ca de uiire. Rorota, 23 Sept 1945,

1 & genitalia [UCLA].

Inini: Saiil, 3 Mar 1944, 1 P, 2 L [UCLA].

GUYANA. Essequibo: Essequibo River, Moraballi Creek, 30 Oct 1929,

1? [BMNH]. Omai, K.S. Wise, 1c, 12 [USNM]. Rio Mazaruni, 25 June 1936,

1 o [UCLA].

NICARAGUA. Potosi, 22 July 1943, 2 L [UCD], 1 L [USNM].

PANAMA. Bocas del Toro: Almirante, 4 May 1963 (PA315), 13 L [UCLA].

Canal Zone: Alhajuelo, 4 Mar-27 May 1912, A. Busck, 70, 1 2 [USNM],

1o [UCLA]. Arraijan, 18 Aug 1950, 5 L [UCLA]. Barro Colorado Island, 23

Dec 1928, C.H. Curran, 1 & genitalia [CU]; Feb 1937, 1c’, 19 [UCLA]. Coro-

zal, 1944, 1% [UCLA]. Corozal Dam, 28 June 1943 (KO3-1), 492 [UCLA]; 22

July 1943, 2 L [USNM]; 12 Jan 1944 (KO3-4), 10, 19 [UCLA], 1 L [USNM];

same data (KO3-5), 12 [UCLA]. Empire, 10 June 1922, J.B. Shropshire, 2 9

[USNM]. Fort Randolph, 15 Sept 1920, 40, 122 [USNM]; 15 Sept 1924, 4¢,

22 [USNM]; 12 Sept, 49 [USNM]. Fort Sherman, 1918, L.H. Dunn, 2 2 [USNM].

France Field, 10 Oct, 3 0°, 59 [USNM]. Gamboa, 11-20 Aug 1923, H.G. Dyar

and R.C. Shannon (P 43), 49, 1 P, 1 L [USNM]; 4 Aug 1923, H.G. Dyar and

R.C. Shannon (P 44), 1 2 [USNM]; 10 July 1945, K.E. Frick (242-1), 9 o, 109,

1P, 10L [CAs], te [UCD]; 11 July 1945, Van Doren, 44 ¢, 41 2, 80L [UCLA],

8L [USNM]. Gatun, 4 June 1934, 2L [USNM]; 24 Sept 1964 (PA 711), 2 L[UCLA);

same data (PA 712), 3 lpo’ (712-41, 42, 48), 2 1p (712-43, 44), 3 p? (712-106-

108), 3c, 89, 11 P, 124L [UCLA]; 2 t. [USNM]. Gatun Lake, Cano Saddle, May

1923, R.C. Shannon, 3%, 12 [USNM]; 5 Aug 1923, H.G. Dyar and R.C. Shan-

non, 1 o, 19 [USNM], 12 [UCLA]. Majagual, 26 Nov 1921, J.B. Shropshire,

7, 82 [USNM]; 8 July 1922, J.B. Shropshire, 1 2 [USNM]; 20 Jan 1923, J.B.

Shropshire, 20, 12 [USNM]. Margarita, 20 Jan 1923, J.B. Shropshire, 2 ?

[USNM]. Mindi, 12 July 1922, J.B. Shropshire, 19 [USNM]; 7 Aug 1923, H.G.

Dyar and R.C. Shannon (P 40), 19 [USNM]. Mt. Hope, 9 Aug 1923, H.G. Dyar

and R.C. Shannon (P 49), 1 oc [USNM]. Rio Trinidad, 5 May 1911, A. Busck

(648), 19 [USNM]. Summit, 30 Aug 1941, W.H.W. Komp, 2P, 1L [USNM]. Sum-

mit Road, 30 Aug 1941 (KO3-2), 1c, 22 [UCLA]; 17 Aug 1941, 19 [UCLA]. Tab-

ernilla, 14 Aug 1908, A. H. Jennings, 1 gynandromorph [USNM]. Toro Point, 11-

28 Feb 1922, J.B. Shropshire, 1c, 12 [USNM]. No locality, 31 Dec 1921, J.B.

Shropshire, 1o [USNM]. No locality or date, H.H. Crowell, 12 [UCLA]; 1, 39

[UCLA]; 1 & genitalia [USNM].

Colon: Portobelo, 5 Dec 1963 (PA588), 1 1p? (588-101), 9 L [UCLA]; 1 ¢

[USNM]. _

Darien: Jaque, 3-4 July 1945, 20°, 22 [UCLA]; 18 Dec 1963 (PA 610),

11lpo (610-108), 31p? (610-102, 105, 107), 1L [UCLA]. Jaque River, 3 July

1945, W. H.W. Komp, 1 1p (37) [USNM]; 19 Dec 1963 (PA 612), 1 1p (612-120),

10 L [UCLA]. La Palma, 29 Nov 1966, O.G.W. Berlin (PA 960), 1 1p? (960-

30) [UCLA]; 10 Dec 1966, O.G. W. Berlin and M. Mena (PA 997), 1 1p (997- 10)

[UCLA]. Tuira River, 1 Apr 1958 (GG 50), 10, 1 9 [UCLA].

Panama: Pacora, 15 Dec 1949, S.J. Carpenter, 10, 492 [CAS]. Pacora,

La Lumbadora (Cerro Azul), 9 Fob 1963 (PA 79), 2 ip¢ (79-112, 114), 3 1p?

(79-104, 110, 113), 4 po” (79-101, 108, 109, 111), 5 p2 (79-102, 103, 105-107), 1 P,

0 L [UCLA]; 13 Feb 1963 (PA 88), 2 L [UCLA]; 18 Feb 1963 (PA 109), 2lpo

(109-105, 106), 4 1p? (109-101, 110-112), 1 po (109-107), 1 p@ (109-109), 41p

(109-102-104, 113), 1¢, 11 L [UCLA].

No locality or date: S.J. Carpenter, 12[UCLA]; A.H. Jennings, 2 L[USNM];

A.H. Jennings (390), 1 lpo (390-16), 3 lp? (390-1,9,14), 29, 1 L [USNM];

A.H. Jennings (420), 20°, 19, 1 L [USNM].

eo Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

PERU. Loreto: Iquitos, Mar-Apr 1931, R.C. Shannon, 1 o, 19 [USNM].

TRINIDAD. Arima, 6-10 Sept 1964, A. Guerra (TR 677), 3 L [UCLA].

Mayaro, 5-16 Dec 1963 (TR 1), 41po (1-101-104) [UCLA]; 6-16 Dec 1963 (TR

2), 41po (2-102, 104,110, 111), 2 1p? (2-103, 105) [UCLA]; 3-23 Dec 1963 (TR

6), 4 1po" (6-101, 103-105), 1 1p? (6-102), 6, 59, 11 P, 11 L [UCLA]; 3-30

Dec 1963 (TR 8), 71po (8-105, 108-110, 113, 114,117), 11 lp? (8-101, 104, 106,

107,111, 112,115, 116, 118-120) [UCLA]; 4-30 Dec 1963 (TR 9), 41lpc (9-104,

105,109,129), 4 1p? (9-106-108, 128), 707, 119, 18P, 17L [UCLA]; 5-30 Dec

1963 (TR 10), 20, 19, 3 P, 2L [UCLA]; 7-30 Dec 1963 (TR 11), 3 1lpo (11-

110, 112,117), 5 1pQ (11-111, 113-116), 9 L [UCLA]; 4-13 Jan 1964 (TR 17),

1 lpo& (17-119), 2 1p? (17-109, 110) [UCLA]; 1-13 Jan 1964 (TR 41), 1 1p (41-

114) [UCLA]; 6-9 Dec 1963 (TR 62), 1 lp (62-126), 21p9 (62-127, 128) [UCLA];

same data (TR 663), 4c, 49 [UCLA]. Saint Anns, 6-7 June 1938, E. Dever-—

teuil, 20°, 29 [BMNH]. Sangre Grande, 10 Sept-1 Oct 1964, A. Guerra (TR

745), 1 [UCLA]; 17 Dec 1964, F. Powdhar (TR 882), 6 L[UCLA]. Sangre

Grande, Tamana Mt., 2-19 June 1964, A. Guerra (TR 500), 1 1p? (500-131)

[UCLA]. Tabaquite, 1914, J.R. Dickson, 1%, 19, 1 L [BMNH]. No locality

or date, F.W- Urich, 4¢, 2°, 21 JUSNM|. :

VENEZUELA. Aragua: Turiamo, 12 Sept 1944, 1 o [UCLA].

: Guaruries, Sept 1943, P.J. Anduze, 1% [USNM].

NO LOCALITY. 17 May 1943, T.H.G. Aitken, 10, 19 [USNM]; 18 Jan

1935, 1 L [USNM]; 18 Mar 1935, 30’, 3 9 [UCLA]. |

NO DATA. (39C), 6 L [USNM]; (575), 20° [UCLA]; (182-4), 1 L [CAS];

(745), 1c genitalia [UCLA]; 2 o (11464, 12690), 2 9 (7587, 12593) [UCLA]; 5 9

[UCLA]; 1 o genitalia [USNM].

Additional records from the literature. BRAZIL. Maranhao: S4o Luis

(Cerqueira, 1961:143).

Minas Gerais: Belo Horizonte, Coronel Fabriciano, Januaria and Monte

Carmelo (Maciel, 1962:489). The material from some of these localities may

possibly be sampaioi.

ECUADOR. Guayas: (Levi-Castillo, 1953:37).

Los Rios: (Levi-Castillo, 1953:37).

PANAMA. Chiriqui: L (Galindo, Carpenter and Trapido, 1955:159, 161).

Cocle: Tucue, A (Trapido, Galindo and Carpenter, 1955:535).

SURINAM. Brokopondo: Kabel Station, L (Bonne and Bonne-Wepster, 1925:

490-494). 7

Marowijne: Moengo, L (Bonne and Bonne-Wepster, 1925:490-494).

VENEZUELA. Merida: Tovar (Anduze, 1947:359-360).

Miranda: (Cova-Garcia, Sutil and Rausseo, 1966b:51-52, 117, 143, 371).

Tachira: Ayacucho and Uribante (Anduze, 1947:359-360).

9. Orthopodomyia sampaioi Lima

: Fig. 18

1935. Orthopodomyia (Orthopodomyia) sampaio Lima, 1935:176-178. *TYPE:

lectotype © (1861) with genitalia in capillary tube, Tijuca, Rio de Janei-

ro, Brazil, adult (?) found in a treehole, June 1934, P.C. Sampaio;

PRESENT SELECTION [IOC, vial 941]. Stated to be probably only a sub-

species of fascipes by Lane (1953:623).

Orthopodomyia sampaioi of Lane (1939:97); Knight.and Mattingly (1950:2); Lane

(1953:623); Stone, Knight and Starcke (1959:124).

Zavortink: Genus Orthopodomyia } 83

Orthopodomyia fascipes of Pinto (1932:295-296); Lane (1936:131; 1937:128).

FEMALE. Wing: 4.63 mm. Proboscis: 3.11 mm. Forefemur: 3.03 mm.

Abdomen: about 3.1 mm. Head: decumbent scales white and yellowish mesal-

ly, mostly white and yellowish laterally, erect scales yellowish mesally, large-

ly light to dark brown laterally; posterior dorsolateral, lateral and ventral

Scales white; palpus 0.36 of proboscis; torus and flagellar scales white. Tho-

rax: integument of mesonotum reddish-brown to chestnut-brown with prescutal

and lower supraalar areas and prescutellar space darker, scutellum lighter

than mesonotum, postnotum reddish-brown to dark brown except for lighter lat-

eral edge, pleuron reddish-brown to dark brown dorsoanteriorly, lighter ven-

trally and posteriorly; acrostichal scale band with large white or cream-colored

scales in more or less complete row throughout length of band, large brown

scales at caudal end, remaining scales large and small, yellow; dorsocentral

scale band with small white and brown scales at anterior promontory, large

white or cream-colored scales in a more or less complete row from level of

posterior fossal area to caudal end, remaining scales large and small, mostly

yellow; lateral prescutellar scales silvery-white posteriorly; lateral prescutal

scales brownish and/or whitish anteriorly and posteriorly or yellowish poste-

_riorly; fossal scales silvery-white and brownish, sometimes some yellowish;

supraalar scales silvery-white anteriorly, in ventral projection and among

bristles, remainder white and yellowish or brownish; scutellar scales white to

silvery-white; pleural scales creamy-white. Legs: forecoxal scales all white

or afew brown, midcoxal scales white above, brown and white below, hindcoxal

scales white above, brown or brown and white below; forefemur largely brown-

scaled with white and yellowish scales in an irregular more or less complete

ring about 0.7-0.8 distance from base and scattered, the scattered light scales

sparse basally, more numerous basad of ring; midfemur like forefemur except

that posterior surface has pale basal streak; hindfemur with anterior surface

largely white- to cream-scaled, brown scales in basal streak, irregular dorsal

patches and scattered, posterior surface entirely white- to cream-scaled in

basal 0.5-0.6, remainder largely light brown-scaled with scattered cream or

yellowish scales; tibiae largely brown-scaled, cream scales at apex, scattered,

and concentrated on posterior surface of fore- and midtibiae and ventral and

anterior surfaces of hindtibia; tarsi largely brown- or black-scaled, segment >

1 with white to cream-colored scales forming narrow basal ring on fore- and

midtarsi, small dorsal basal patch on hindtarsus and usually scattered; fore-

tarsal segment I dark-scaled at apex and segments 2-5 entirely dark-scaled or

white scales present at apex of segments 1-4 and base of segment 3; midtarsus

with white to cream-colored scales covering apex of segment 1, ends of segment

2, ends or entire surface of segment 3 and all of segments 4 and 5; hindtarsal

segment 1 with or without a very few white scales at apex, segment 2 with nar-

row to moderately broad white-scaled apical ring, segment 3 entirely white-

scaled, segment 4 with white scales in dorsal patch or narrow ring at ends,

segment 5 white-scaled dorsally. Wing: light scales almost entirely white,

rarely some scattered ones dingy; veins R9.3 and M without distinct patch over

furcation; vein R445 with an apical light patch. Haltere: integument of stem

whitish, knob beige; scales on stem white, those on knob transparent to brown.

Abdomen: sternites II-VII dark-scaled with large basolateral light-scaled patch

and sometimes poorly developed light basal band, light patch on distal segments

apparently larger than in fascipes.

MALE. Similar to female. Palpus: 0.93 of proboscis. Antenna: flagellar

scales white. | .

84 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

MALE GENITALIA (fig.18). Segment VUI: posterior margin of tergite

slightly rounded mesally. Sidepiece: very broad basally, sharply constricted

distad of basal mesal lobe; basal mesal lobe with 6-8 long fine bristles which

are not separated from strong blunt bristles by a gap; mesal and tergal sur-

faces near apex of sidepiece with patch of specialized bristles, individual bris-

tles directed tergally or tergomesally and curved mesally at apex; tergomesal

surface distad of basal mesal lobe with poorly differentiated short, flat curved

bristles. Aedeagus: broader distad of middle; each half of sternal surface with

sclerotized tooth along mesal edge; without projection between or below ventral

parameres. Proctiger: paraproct sclerotization with 2 apical teeth; cercal se-

tae 2-5. |

PUPA. Unknown.

LARVA. Lima (1935:178) states that the larva resembles fascipes. I have

examined 2 fourth instar larvae from Tijuca, Rio de Janeiro, Brazil, which are

presumably this species. Although both these larvae are in very poor condition

and cannot be critically studied, it has been noted that they differ from fascipes

as follows: (1) head capsule and antenna without spicules, (2) hair 1-A weakly

developed and only 2-4b, (3) hair 1-M single, (4) 1 branch of hair 2-K much

elongated, and (5) ventral brush with only 7 pairs of hairs.

BIONOMICS. Larvae have been collected in treeholes. Nothing is known

of the habits of the adults.

SYSTEMATICS. Sampaioi is evidently a relict species restricted to south-

ern Brazil. The characteristics by which the adults and male genitalia differ

from fascipes are indicated in the keys and discussed in the systematics sec-

tion of fascipes.

The larva of sampaioi is so incompletely known that it has not been included

in the keys. The specimens examined differ from fascipes by the features indi-

cated in the description. The affinities of the Thomasina section may become

more obvious when the immature stages of sampaioi are completely known.

DISTRIBUTION. Goias, south to Parana and Guanabara. Material Exam-

ined: 13 specimens; 50°, 59, 3 L.

BRAZIL. Goias: Anapolis, Mar 1936, 1 o% [USNM].

Guanabara: Rio de Janeiro, Tijuca, June 1934, P.C. Sampaio, © lectotype

[IOC]; 3 L [USNM].

Parana: Londrina, 2 o° [USNM], 1 2 [BMNH].

Sao Paulo: Avare, 1 [CU]. Serra do Cubat&io, Dec 1929, O. Castro, 1 9

[IOC].

NO DATA. (12226), 29 [UCLA]; 1 2 (12690) [UCLA].

Additional records from the literature. BRAZIL: Minas Gerais: vide fas-

cipes.

FOLICOLAE SECTION

10. Orthopodomyia phyllozoa (Dyar and Knab)

Figs. 2,19, 20 —

1907. Mansonia phyllozoa Dyar and Knab, 1907:199. *TYPE: holotype o (177.2),

native village near Tabernilla, Canal Zone, Panama, reared from a larva

found in a bromeliad leaf axil, 25 June 1907, A. Busck [USNM, 10864].

Orthopodomyia phyllozoa of Howard, Dyar and Knab (1917:879-882); Dyar (1923a:

183); Bonne and Bonne-Wepster (1925:489); Dyar (1925:179; 1928:395-396);

Zavortink: Genus Orthopodomyia 85

Edwards (1932:107, 108; 1939:121); Lane (1939:97); Kumm, Komp and Ruiz

(1940:405, 417,418); Anduze (1941:12; 1947:360); Knight and Mattingly (1950:

1); Lane (1953:626-628); Stone, Knight and Starcke (1959:123); Cova-Garcia,

Sutil and Rausseo (1966a:53; 1966b:52, 143, 371). :

Bancroftia phyllozoa of Picado (1913:353).

_ Mansonia phyllozoa of Busck (1908:60); Theobald (1910b:446, 619).

FEMALE (fig. 2). Wing: 2.59 mm. Proboscis: 1.32 mm. Forefemur:

1.62 mm. Abdomen: about 2.0 mm. Head: integument of head capsule, cly-

peus and flagellum light to dark brown, torus brown to dark brown dorsally,

lighter brown or yellowish ventrally; eyes separated above antennae by a row

of silvery-white decumbent scales; frontal bristles amber to brown; orbital

bristles amber to brown, 1 or 2 lateral pairs and 1 or 2 mesal pairs; ocular

border absent; vertex and occiput with numerous narrow curved decumbent

scales and numerous erect scales, decumbent scales broader and denser along

edge of eye, yellow laterally, silvery-white mesally, remaining decumbent |

scales mostly bright yellow or yellow and silvery-white, sometimes a few brown

or cream, erect scales mostly yellow, but usually a few tan or brown; dorso-

lateral scales brown anteriorly, yellow posteriorly; suborbital bristles pale;

lateral and ventral scales mostly yellow; clypeus more or less triangular, bare;

labium considerably swollen apically, shaft bristles absent, mostly brown-

scaled, usually with white and/or yellow scales forming dorsal patch 0. 67-0. 75

distance from base and dorsal preapical patch and with few to very numerous

scattered yellow scales; palpus usually coiled in dried specimens, 0. 54 of pro-

_ boscis, 4-segmented, segment 4 short, largely brown-scaled, yellow scales

usually in patch over joint between segments 2-3 and scattered throughout en-

tire length, silvery-white scales covering segment 4; torus with mesal patch

of silvery flat or hair-like scales; flagellar segment 1 swollen and with tuft of

brown scales; dorsal flagellar bristles about 2.5-3.0 length of their segment

on proximal segments, ventral bristles absent on segment 1 and usually on seg-

ments 2-3 or 4. Thorax: integument of mesonotum and pleuron light rusty |

brown to dark brown except for lighter area on mesonotum near prescutal scales,

scutellum lighter, postnotum usually darker; paratergite narrow, bare; poste-

rior dorsocentral bristles usually not in double row; humeral, lateral prescutal

and posterior fossal bristles not numerous enough to completely enclose fossa;

fossal bristles absent; mesonotal bristles dark brown, parascutellar and scu-

tellar bristles amber; mesonotal ornamentation consisting of mostly brown and

silvery-white scales in longitudinal bands; brown scales arranged as follows:

(1) in band accompanying the acrostichal bristles, and (2) in band accompanying

the dorsocentral bristles; larger silvery-white or rarely yellow-tinged scales |

arranged as follows: (1) acrostichal scales in small patch on anterior promon-

tory, (2) lateral prescutal scales extending from anterior promontory to scutal

angle along edge of mesonotum, (3) posterior fossal scales extending from scu-

tal angle to dorsocentral scales anterior of scutal angle, (4) lateral prescutel-

lar scales larger than other mesonotal scales, accompanying lateral prescutel-

lar bristles and extending caudad to margin of scutellum, and (5) supraalar

scales extending from caudad of scutal suture to cephalad of parascutellum

mesad of supraalar bristles; additional silvery-white scales sometimes later-

ad of posterior dorsocentral brown scales; golden supraalar scales present

mesad of paratergite; parascutellum normally without scales; lateral and mid-

scutellar lobes each with an extensive patch of large silvery-white scales which

extend over the caudal margin of the scutellum; ppn bristles brown, remaining

pleural bristles amber; upper anterior apn bristles 3-6 strongly developed,

86 Contrib. Amer. Ent. inst., vol. 3, no.°2, 1968

2-5 weakly to moderately developed, lower apn bristles 1 or 2, weakly to strong-

ly developed; ppl bristles 3 or 4 strongly developed, 1-3 weakly to moderately

developed; pra usually with 1 bristle (0-2); upper stp bristles 1, strongly devel-

oped, posterior stp bristles 2-4, moderately to strongly developed, lower stp

bristles 3 or 4, weakly to strongly developed; upper posterior mep bristles 1-4,

moderately developed, anterior mep bristles absent; pleural scaling consisting

of patches and lines of curved and flat silvery-white, yellowish and yellow scales;

apn with yellowish scales in a diagonal row between upper and lower bristles;

ppn with silvery-white to yellowish scales along posterior margin below bris-

tles; psp and pra without scales; ppl with yellowish to yellow scales in patch

around bristles; pst with yellowish scales only near ppl; pcx, ssp and hypostig-

- ial scales absent; stp with 2 horizontal rows of scales, 1 above and 1 below up-

per stp bristle, upper row with silvery-white to yellowish scales, lower row

with yellowish to yellow scales, the 2 rows joined anterior to bristle by vertical

row of yellowish to yellow scales, and upper row with vertical extension of sil-

very-white to yellow scales to pra, lower portion of stp with semierect and flat

scales in vertical area along posterior margin; mep with semierect silvery-

white scales in an upper patch anterior to mep bristles and semierect and flat

silvery-white to yellowish scales in a small patch near anterior edge opposite

lower stp horizontal scale row; a few additional large brownish or transparent

scales may be scattered over the pleuron. Legs: integument of coxae same

color as pleuron; coxal bristles amber and/or brown; forecoxa with broad flat

yellowish scales above and behind uppermost bristles and with smaller brown

or brown and yellow scales on anterior surface and among bristles, mid- and

hindcoxae with broad flat scales on anterior surface, those on midcoxa yellow

above, brown below, those on hindcoxa brown; trochanters with brown or brown

and yellowish flat scales on anterior, inner and posterior surfaces; femora

with mixed brown and yellow scales except for small white knee spots, largely

brown-scaled preapical ring on fore- and midfemora and yellowish-scaling in

basal 0.3-0.6 of posterior surface of mid- and hindfemora, the scattered yel-

low scales concentrated in basal portion of anterior surface of forefemur and

in basal and apical portions of anterior surface of hindfemur; tibiae brown- or

black-scaled with numerous irregular yellowish-scaled rings or patches and

white- or yellowish-scaling at apex; hindtarsal segment 1 longer than hindtibia;

tarsi largely brown- or black-scaled, segment 1 with cream-colored or yellow-

ish scales forming narrow basal ring on fore- and midtarsi, dorsal basal patch

on hindtarsus and sometimes poorly developed irregular rings or patches on all

tarsi; foretarsal segment 1 dark-scaled at apex, segments 2-09 entirely dark-

scaled or with white or cream-colored scales in dorsal patch at base of seg-

ment 2; midtarsus with white or cream-colored scales forming complete or in-

complete narrow ring or small dorsal patch at apex of segment 1, complete

narrow ring at base of segment 2, small dorsal patch or incomplete ring at

base of segment 3, and sometimes dorsal patch over segments 4 and 5; hind-

tarsus with white or cream-colored scales forming broad ring at apex of seg-

ment 1 and base of segments 2 and 3, narrower complete or incomplete ring

or dorsal patch at apex of segment 2, narrow incomplete ring or dorsal patch

at apex of segment 3, dorsal basal patch on segment 4 and basal or more ex-

tensive dorsal patch on segment 5; hindclaws smaller than fore- and midclaws.

Wing: remigial bristles absent; vein Rg 1.36 length of vein R2+3; vein Mj.9

1.18 length of vein M distad of crossvein m-cu; plical area with or without a

few dark scales; alula scales moderately broad; upper calypter with scale-like

bristles; scales symmetrical, moderately broad, mostly spreading; wing large-

ly brown- or black-scaled with white to yellowish scales in quite variable

Zavortink: Genus Orthopodomyia 87

patches, lines and speckles, as follows: (1) costal vein with basal, accessory

sectoral, subcostal and apical light patches and usually with presectoral and

accessory subcostal light patches, (2) vein Sc with accessory sectoral and sub-

costal light patches and usually with presectoral light patch, (3) vein R light-

scaled from base to or beyond level of distal end of presectoral costal patch,

(4) vein R1 with accessory sectoral, subcostal and apical light patches and usu-

ally with accessory subcostal light patch, (5) veins Rg and R445 with light patch

at base, (6) vein Ro9,3 with light patch opposite base of vein R4+5, (7) veins

R2+3, M and sometimes Cu with light patch over furcation, (8) veins M and Cu

with light patches distad of crossveins, (9) some or all of veins Ro, Rg, R4+5;

M1+2, M344, Cug and 1A with small apical light patch, (10) veins M and Cu

usually with scattered light scales in basal portion, sometimes with distinct

basal light patch, (11) vein 1A usually with scattered light scales, sometimes

with distinct central light patch, and (12) vein Cuy sometimes with scattered

light scales or central or more extensive light patch. Haltere: integument

beige, Scales whitish to yellowish. Abdomen: integument of tergites and distal

sternites brown to dark brown, proximal sternites lighter, even transparent;

laterotergite bare; tergite 1 with dark brown or dark brown and yellow scales

in fan and sometimes with a few other scattered scales; remaining segments

with dark scales brown or black, light scales whitish to yellowish; tergites

largely dark-scaled, segments II-VI or VII with light scales in a more or less

diagonal patch starting basolaterally, segment VIII or segments VII and VIII

with light scales in smaller basolateral patch; proximal tergites sometimes

with a few dorsobasal or other light scales; sternites largely dark-scaled, seg-

ments II and III with large basolateral or lateral light patch, segments IV-VIII

with light scales in smaller basolateral patch or poorly developed basal band.

FEMALE GENITALIA. Tergite IX without dorsolateral setae.

MALE. Similar to female. Labium: not more strongly swollen than in fe-

males, yellow to white scales usually in narrow ring distad of level of distal end

of palpal segment 2, in dorsal preapical patch and scattered. Palpus: 0.88 of

proboscis, segment 5 very short, segment 3 normally without strong apical

bristles; largely brown-scaled, yellow scales at joint between segments 1-2

and 2-3 and scattered on segments 2 and 3, silvery-white scales at joint be-

tween segments 3-4 and covering all of segment 5. Antenna: integument of fla-

gellum dark brown except for lighter rings basad of bristles on shorter seg-

ments; torus greatly enlarged; flagellum strongly plumose; segment 13 about

1.5 length of segment 12; flagellar segment 1 with tuft of brown scales; bristles

of flagellar whorls about 0.4-0.6 length of entire flagellum on proximal seg-

ments, about 0.4-0.8 length of segment 13 on that segment. Legs: anterior

fore- and midclaws with basal tooth too small to be seen on pinned specimens,

posterior fore- and midclaws simple. Abdomen: light scaling on tergites some-

times more extensive than in female; sidepiece with light and dark scales.

MALE GENITALIA (fig. 19). Segment VIII: posterior margin of tergite with

distinct lobe; lobe as long as broad to distinctly longer than broad, sides paral-

lel, diverging or converging apically, apex truncate or more often emarginate.

Segment IX: tergite usually without setae dorsolaterally, sometimes with 1 fine

seta on each side. Sidepiece: long conical; basal mesal lobe with 3-5 strong

terete apical bristles which are slightly bent mesally at apex and usually with

1 or 2 finer apical bristles; mesal surface membranous distad of basal mesal

lobe; without specialized mesal bristles but usually with 1-3 specialized, curved

tergomesal bristles distad of basal mesal lobe. Spiniform: 1, apex slightly flat-

tened. Aedeagus: moderately sclerotized; apex with 2-7 large teeth on each

side; base with short projection between ventral parameres and with 2 strongly

88 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

sclerotized, tergally-directed processes; sternal surface strongly sclerotized

along mesal edge, without teeth. Proctiger: paraproct sclerotization with 1-3

apical teeth; cercal setae 1-3.

PUPA (fig.19). Abdomen: 3.11 mm. Trumpet: 0.46 mm. Paddle: 0.59

mm. Cephalothorax: uniformly very lightly pigmented or with upper posterior

portion of wing case and portion of mesonotum caudad of trumpet slightly dark-

er; integumentary sculpturing weak, reticulate; smaller hairs concolorous with

integument, larger hairs darker; hairs 1, 2,4, 7-C moderately and more or less

equally developed, usually 2-4b (2-5); 3-C weakly developed, 2-4b, and much

smaller than 1 or 2-C; 5-C usually 3, 4b (2-5), very strongly developed, at

least 2 times length of 4-C; 8,9-C moderately developed, usually single; 6-C

cephalad of 7-C. Trumpet: uniformly lightly pigmented and only slightly dark-

er than darkest portion of cephalothorax; distal portion gradually widened from

base, rather narrow; pinna small. Metanotum: lightly pigmented with haltere

cases lighter; integumentary sculpturing weak to moderate, reticulate; never

more than 3 pairs of hairs developed. Abdomen: very lightly pigmented, with

distal segments lighter; anterior portion of proximal segments darker; ante-

rior segments with moderately developed imbricate integumentary sculpturing,

posterior segments with weaker sculpturing; smaller hairs concolorous with

integument, larger ones darker; hair 1-II dendritic, usually with about 15-30

(9-41) ultimate branches; 2-II cephalad of 3-II; 5-IV-VI usually 3b (2-4) and

with 1 branch very long, reaching nearly to or extending beyond alveolus of hair

4 of second following segment; 1-VI, VII laterad of hair 2 of corresponding seg-

ment; 9-VII weakly to moderately developed, usually 3-6b, longest branch ex-

tending about 0.4-0.7 distance between alveoli of hairs 9-VII and 9-VIII; 9-VIII

strongly developed, usually 9-13b, longest branches reaching nearly to or be-

yond apex of paddle. Terminal Segments: male genital lobe projecting consid-

erably beyond median caudal lobe. Paddle: very lightly pigmented, with mid-

rib somewhat darker; longer than broad, more or less elliptical in outline,

never angulate; external buttress strongly convex laterally, with minute mar-

ginal spicules; midrib usually almost straight; entire surface of paddle covered

with minute spicules.

LARVA (fig. 20). Head: 0.79 mm. Siphon: 1.06 mm. Anal Saddle: 0.30

mm. Head: distinctly broader than long; integument with minute spicules in

posterior dorsal portion; largely straw-colored, slightly lighter around imagi-

nal eye, darker caudally, collar brown; labium moderately long; mental plate

brown, usually with 8,9 (7-10) teeth on each side; hair 2-C absent. Antenna:

moderately long; integument with small spicules; uniformly straw-colored.

Thorax: epidermal pigment present, weakly to moderately developed, purple

in color; spicules not conspicuous; hairs moderately pigmented; hair 8-P weak-

ly developed; 1-M, T moderately developed, multiple; 4-M always double or

multiple. Abdomen: segment VI without sclerotized plate, segment VII with

large quadrangular sclerotized plate; hairs moderately pigmented; hairs 1-I- |

IV moderately developed, multiple; 2-I, II mesad of hair 1 of corresponding

segment; 6-I, II multiple, 6-III-VI single; 13-I longer than 10-I; 5-II-VI mod-

erately developed, usually 3b; 13-II-V moderately developed, multiple; 2-VI,

VII mesad of hair 1 of corresponding segment. Segment VIII: large sclerotized

plate present; comb scales in 2 rows; anterior comb scales usually 16-24 (12-

26), posterior comb scales usually 2-4 (1-6); anterior and posterior comb

scales of the same size and shape, with a medium-sized apical spine flanked

on either side by 1-3 smaller spines which merge basally with fringe. Siphon:

base brown, remainder uniformly straw-colored or apical portion slightly dark-

er; index 4.5-9.0; hair 1-S located 0.35-0.50 from base of siphon, longest

Zavortink: Genus Orthopodomyia | 89

branches extending 0.55-0.90 distance from alveolus to apex of siphon; moder-

ately to strongly developed imbricate integumentary sculpturing present basal-

ly. Anal Segment: saddle complete, base and apical portion usually tan, re-

- mainder uniformly light straw-colored or with darker areas dorsally; sclero-

tized band or incomplete ring present at base of saddle; integumentary sculp-

turing weaker than on siphon; ventral brush usually with 7 pairs of hairs, the

more caudal hairs usually with about 10-12 branches; gills moderately to very

long, dorsal pair about 0.95-1.5 as long as ventral pair, 1.5-5.5 length of anal

saddle and 0.4-2.1 length of siphon.

BIONOMICS. The immature stages are found in the leaf axils of bromeli-

ads and the spathes of heliconias. Nothing is known of the habits of the adults.

SYSTEMATICS. Phyllozoa is so distinct from other Orthopodomyia in all

stages that it is worthy of recognition as a separate section. The most distinc-

tive features are the ornamentation of the adult thorax, the shape and dentition

of the aedeagus, the development of pupal hairs 1-7-C and 9-VII, VII and shape

of the larval head and comb scales. :

Larvae of phyllozoa collected in heliconias differ quite strikingly from those

collected in bromeliads by their shorter siphons and longer anal gills. In the

heliconia form the siphon index varies from 4.5-5.5 and the gills are usually

1.3 to 2.1 times the length of the siphon; in the bromeliad form the siphon in-

dex varies from 6.0 to 9.0 and the gills are 0.4-0.6 the length of the siphon.

The only apparent difference in the chaetotaxy of the 2 forms is that hair 8-P

usually has a greater number of branches in the larvae from heliconias. Pupae

from the 2 habitats are very similar, but those from heliconias differ by slight-

ly longer paddles and fewer branches in hair 5-C. Adults bred from the 2 lar-

val forms are apparently indistinguishable and there are no differences in the

male genitalia. For the present I am considering the heliconia and bromeliad

forms to be conspecific. It is possible, however, that study of living material

may show that 2 distinct species are involved.

Although extremely hairy forms of pupae and larvae are unknown, pupae

with hairs 1,3,'7,10-12-C, 6, 7-I and 6-II longer and stronger than customary

and larvae with the branches of the moderately developed multiple hairs of the

thorax and abdomen stouter and more numerous than usual have been seen in

collections from both heliconias and bromeliads.

Adults of phyllozoa exhibit considerable variation, particularly in ornamen-

tation of the torus, proboscis, wing and first tarsal segments. The torus may

have a conspicuous patch of silvery-white broad scales or an inconspicuous

patch of silvery-white hair-like scales. The proboscis usually has 2 dorsal

light patches and scattered light scales, but either or both patches may be ab-

sent or the scattered light scales may be so numerous as to obscure them. The

wing may or may not have presectoral and accessory subcostal light patches

and the light-scaling on veins M, Cu and 1A is quite variable in extent. The

first tarsal segments vary from being nearly entirely dark-scaled to eaihates

conspicuous light patches or rings.

Phyllozoa appears to be an ancient species which has escaped being re-

placed by more recent ones by occupying different habitats. It combines char-

acteristics found in other groups of Orthopodomyia and cannot be definitely al-_

lied with any. Edwards (1932:107; 1939:121) placed phyllozoa in Group A (Or-

thopodomyia) because of its wing and leg ornamentation but Knight and Matting-

ly (1950:1) moved it to Group B (Bancroftia) on the basis of the male genitalia

and mesonotal ornamentation. Consideration of the immature stages does indi-

cate a distant kinship with the Oriental Albipes group; in particular, the larval

chaetotaxy of wilsoni and phyllozoa is similar.

90 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

Phyllozoa is sympatric with 3 other species of Orthopodomyia: albicosta,

fascipes and kummi. As discussed more fully in the systematics section of fas-

cipes, some of the characteristics of that species may have been acquired from

phyllozoa by introgression.

DISTRIBUTION. Nicaragua and north-central Venezuela, south to central

Colombia. Material Examined: 944 specimens; 89 c, 130 2, 1 isolated © geni-

talia, 187 P, 537 L; 181 individual rearings (103 larval, 53 pupal, 25 incom-

plete). —

COLOMBIA. Meta: Restrepo, 1935, E. Osorno, 1 ¢ [CU].

Valle del Cauca: Buenaventura, Rio Raposo, 20 Oct 1964, V.H. Lee (COL

11), Pipe (11420), tip? (11-29),:1-9,.1 P, 3: (UCLA); 8 Dee 1964, Vv. H.

Lee (COL 33), 31po% (33-10, 14,15), 1 po (33-103), 1 p? (33-100), 10, 29,

19 L [UCLA]; 8 Feb 1965, V.H. Lee (COL 50), 2 lp (50-20, 23), 71p? (50-

21, 22, 25-29), 1 p? (50-24), 22 L [UCLA]; 10 Feb 1965, V.H. Lee (COL 55),

1 po (55-100) [UCLA]; 23 Feb1965, V.H. Lee (COL 57), 2 lpo (57-13, 19), lpo

(57-107), 2 p? (57-100, 106), 1 lp (57-12), 3¢, 19; 3 P, 6: |UCLA|; 2 Mar

1965,'V¥.H. Lee (COL 58); 2 1pc (58-37, 38), 1:ip? (58-32), 1 po’ (58-110),

110 (58-31), 1 lp (58-30), 10 L [UCLA].

No locality or date: E. Osorno, 1 2 [USNM]. |

COSTA RICA. Cervantes, 9 Nov 1962, J.N. Belkin, C. L. Hogue and W. A.

Powder (CR 13), 2 lp? (13-205, 206) [UCLA]. Orosi, 7 Dec 1962, C.L. Hogue

and W.A. Powder (CR 83), 4lpo (83-201, 203, 207, 209), 51p2 (83-202, 205,

206, 212, 213), 1 1p (83-211), 35L [UCLA]; 4 Jan 1937, 19 [USNM]; 6 Jan 1938,

1 9 [BMNH]; Nov, C. Picado, 20, 3 2 [USNM]; 6 [BMNH]. — |

NICARAGUA. Zelaya, Recreo, 26 June 1954, K. Neiland (N14), 3 L[UCLA];

27 June 1954, K. Neiland (NI 5), 2 L [UCLA].

PANAMA. Bocas del Toro: Almirante, 2 May 1963 (PA 297), 1 po (297-

102), 1 p? (297-103), 7 L [UCLA]. Almirante, Nigua Creek, 27 Apr 1963 (PA

258), 1 p? (258-101) [UCLA]; 28 Apr 1963 (PA 276), 1 L [UCLA]; 29 Apr 1963

_ (PA 279), 1 L [UCLA]; 6 May 1963 (PA 325), 4 L [UCLA]. Big Creek, 10 Apr

1964 (PA 655), 7L [UCLA]. Chiriquisito, El Guabo, 16 Apr 1963 (PA 209),

3 L [UCLA]; same data (PA 212), 1 L [UCLA]. Chiriquisito, Kaysan, 16 Apr

1963 (PA 215), 1 L [UCLA]; 18 Apr 1963 (PA 229), 3 L [UCLA].

Canal Zone: Gatun, 24 Sept 1964 (PA711), 1 lpo (711-20), 7 lp? (711-21-

24,27, 29,30), 3 po (711-105, 112,114), 5 p? (711-26, 100, 103, 106, 108), 3 1p

(711-25, 28, 33), 76L [UCLA]; 4 June 1934, 10°, 19 [UCLA], 2 L [USNM]; 2L

[USNM]. Gatun Lake, Cano Saddle, 7 Aug-25 Oct 1923, H.G. Dyar and R.C.

Shannon (P 67), 20°, 32 [USNM]. Madden Road Forest Preserve, Cruces

Trail, 17 Sept 1964 (PA 709), 1 lp? (709-19) [UCLA]. Tabernilla, A. Busck

(177.2), & holotype [USNM]. No locality or date, A.H. Jennings (416), 1¢

[USNM]; 10’, 1@ [UCLA].

Cocle: El Valle, 19 Apr 1942, G.B. Fairchild, 1 o [UCLA], 1 L [USNM].

Colon: Buena Vista, 24 Sept 1964 (PA 710), 5 1p? (710-23, 24, 26-28), 3L .

[UCLA]. Portobelo, 8-20 Aug 1923, H.G. Dyar and R.C. Shannon (P 79), 1,

10 9 [USNM], 1 o [BMNH]; 10 Feb 1909, A.H. Jennings (480), 1 o [USNM]; 16

Feb 1909, A.H. Jennings (497), 19 [USNM]; 4 Dec 1963 (PA 579), 4 1p? (579-

125, 127, 132, 136), 55L [UCLA]; 9 Dec 1963 (PA 599), 2 po (599-106, 110), 1P,

51 L [UCLA]. Portobelo, Caldera Island, 24 Jan 1908, A.H. Jennings, 1 lp?

(194.1) [USNM]. Santa Rosa, Juan Mina, 17 Jan 1963 (PA3), 2 lpo (3-145, 160),

5 1p? (3-147, 148, 156, 157,161), 2 po’ (3-112, 126), 4 L [UCLA]; 22 Jan 1963

(PA 33), 1 p? (33-103), 1 1p (33-108), 1L [UCLA]; 29 Jan 1963 (PA49), 21pc

(49-108, 112), 2 1p? (49-104, 118), 2 p? (49-116, 132), 1 1p (49-137), 4L [UCLA];

30 Jan 1963 (PA 52), 21lpo’ (52-103, 127), 41pQ (52-106, 116, 121,126), 3 po

Zavortink: Genus Orthopodomyia 91

(52-107, 111,125), 1 p@ (52-123), 4L [UCLA]. Santa Rosa, Rio Chilibre, 21

Jan 1963 (PA 28), 1 1p (28-106), 1L [UCLA]; same data (PA 29), 1 1p (29-109),

Hy [UC bak)

Darien: Jaque, 6 July 1945, 19 [USNM]. Jaque, Rio Jaque, 19 Dec 1963

(PA 613), 1 lp? (613-109) [UCLA]. La Palma, 28 Nov 1966, O.G.W. Berlin

(PA 955), 1 lp? (955-21) [UCLA]; 1 Dec 1966, O.G.W. Berlin and R. Hinds

(PA 962), 1 1po’ (962-12), 1 p? (962-111) [UCLA]; same data (PA 963), 1 lpo"

(963-20) [UCLA]; same data (PA 964), 2L [UCLA]; same data (PA 966), 1 lpo&

(966-13) [UCLA]; 3 Dec 1966, O.G.W. Berlin and R. Hinds (PA 974), 1 lp¢c

(974-11), 1 lp? (974-10), 1 P, 1 L [UCLA]; 7 Dec 1966, O.G.W. Berlin and

R. Hinds (PA 984), 1 1po (984-50), 4 1p? (984-10, 11, 13, 15), 1 po’ (984-104),

3p? (984-108, 109,111), 2L [UCLA]; same data (PA 985), 1 lpo (985-10)

[UCLA]; same data (PA 986), 1 po (986-105) [UCLA]; same data (PA 987),

1 lp? (987-10) [UCLA]. Pucro, Tacarcuna River, 14 June 1963 (PA386), 3 lp&

(386-109, 110, 115), 8 1p? (386-104, 107, 108, 111, 117, 119, 120, 122), 4 po” (386-

101, 102, 106, 113), 1 p@ (386-112), 1 1p (386-123), 15 L [UCLA]; 22 June 1963

(PA 414), 1 lpo (414-101), 1 1p? (414-121), 1 po (414-116), 2 1p (414-118, 120)

[UCLA]; 27 June 1963 (PA 430), 1 1po (430-101), 1 1p (430-103) [UCLA]; 5 July

1963 (PA 435), 1 p? (435-103), 1P [UCLA]. Tuira River, 8 Mar 1958 (GG 89),

1 2 [UCLA]. | |

Panama: Chica, Cerro Campana, 3 Aug 1963 (PA 461), 1 1p? (461-101),

5L [UCLA]; same data (PA 462), 1 po (462-104), 1 1p (462-10), 19, 1 P [UCLA];

5 Aug 1963 (PA 473), 3 lpo’ (473-115, 117, 122), 3 1p9 (473-107, 113, 114), 4 po

(473-102, 110, 123, 124), 3 p? (473-104, 105, 119), 119 (473-106), 8 1p (473-112,

116, 118, 120, 121, 125-127), 2 o', 32 L [UCLA]; 12 Aug 1963 (PA 496), 1 lp¢

(496-105), 1 pot (496-103), 1 p? (496-102), 10°, 12L [UCLA]; 31 Aug 1963 (PA

945), 1 lpo (545-103), 7 L [UCLA]. Coscajal River, 18 Feb 1909, A. H. Jen-

nings (503), 1 o [USNM].

NO LOCALITY OR DATE. A.H. Jennings (157), 30, 2 9 [USNM ].

NO DATA. 1° genitalia [USNM]. :

Additional records from the literature. VENEZUELA. Aragua (Cova-Gar-

cia, Sutil and Rausseo, 1966b:52, 143, 371).

Tachira: Uribante (Anduze, 1947:360).

ORTHOPODOMYIA SECTION

FEMALES. Head: eyes separated above antennae by a row of decumbent

scales which are sometimes accompanied by weakly developed pale or dark

bristles; frontal bristles usually dark; orbital bristles dark, usually 1-3 lat-

eral pairs and 2-4 mesal pairs; ocular border absent; vertex and occiput with

numerous narrow curved decumbent scales which are usually broader and dens-

er at base of frontal bristles and in front of dorsolateral broad scales and very

numerous erect scales; suborbital bristles pale; clypeus an elongate angular

lobe, usually bare; labium slightly to considerably swollen apically, shaft bris-

tles absent; palpus straight, 0.37-0.58 of proboscis, 4- or 5-segmented; inner

half of torus with small scales except for a bare spot mesally and dorsal portion

of inner half with short, fine hairs; flagellar segment 1 not swollen; flagellar

segment 1 with scales; dorsal flagellar bristles about 2.0-3.0 length of their

segment of proximal segments, ventral bristles absent on segment 1 and often

reduced or absent on segments 2-4. Thorax: paratergite moderately broad,

with (flavicosta, flavithorax) or without scales; posterior dorsocentral bristles

sometimes in a double row; humeral, lateral prescutal and posterior fossal

92 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

bristles not numerous enough to completely enclose fossa; fossal bristles ab-

sent; mesonotal ornamentation consisting of light and dark scales of various

sizes and densities almost completely but sparsely covering the scutum, as fol-

lows: (1) acrostichal scales in a broad band accompanying the acrostichal bris-

tles, (2) anterior prescutellar scales in a more or less triangular area anterior

to prescutellar space, (3) dorsocentral scales in a broad band accompanying the

dorsocentral bristles, (4) lateral prescutellar scales accompanying the lateral

prescutellar bristles and extending caudad to margin of scutellum, (5) lateral

prescutal scales extending from anterior promontory near dorsocentral scale

line to scutal angle along edge of mesonotum, small, (6) posterior fossal scales

extending from the scutal angle to the dorsocentral scales anterior of the scutal

suture, small, (7) fossal scales in the area enclosed by the dorsocentral, lat-

eral prescutal and posterior fossal scales, small, (8) supraalar scales extend-

ing from caudad of scutal suture to cephalad of parascutellum among and mesad ~

of supraalar bristles and with anterior and middle extensions ventrad to near —

margin of paratergite; parascutellum normally without scales; lateral and mid-

scutellar lobes each with an extensive patch of large white to silvery-white

scales which extend over the caudal margin of the scutellum; pleural bristles

dark and light; upper anterior apn bristles brownish-amber to brown, 2-6

strongly developed, 0-7 weakly to moderately developed, lower apn bristles

amber, 2-6, weakly to moderately developed; ppn bristles brown; ppl bristles

amber, 2-5 strongly developed, 0-4 weakly to moderately developed; anterior

mep bristles absent; pleural scaling consisting of extensive areas of broad flat,

narrow curved and broad semierect scales; apn with narrow curved scales in

an upper anterior patch and narrow curved or broader and/or flatter scales in

a ventral diagonal row; psp scales normally absent; pra with a few flat or

curved scales on anterior edge and/or below knob; ppl with broad flat and semi-

erect scales in a dense patch around the bristles; entire pst with a dense cov-

ering of broad flat scales; pcx scales absent; ssp scales broad and flat; hypo-

stigial scales present (madrensis) or absent; upper half of stp completely cov-

ered with broad flat scales except for 1-3 small bare areas, lower half with a

vertical row of broad flat and semierect scales; mep with elongate semierect.

white scales in an upper patch anterior to the upper posterior bristles; a few

additional large brownish or transparent scales may be found scattered over

the pleuron, especially the meron and lower stp. Legs: trochanters with white

to silvery-white or slightly yellowish flat scales on anterior, inner and poste-

rior surfaces; femora with small white to dingy knee spots; tibiae white- or

cream-scaled apically; hindtarsal segment 1 longer than hindtibia; hindclaws

slightly to considerably smaller than fore- and midclaws. Wing: remigial bris-

tles absent; plical area usually without scales, sometimes with a few light or

dark scales; alula scales moderately broad; upper calypter with bristles.

FEMALE GENITALIA. Tergite IX with dorsolateral setae.

MALES. Similar to females. Labium: not more strongly swollen than in ©

females. Palpus: 0.75-1.00 of proboscis, segment 5 short. Antenna: integu-

ment of flagellum moderate or dark except (usually) for lighter bands basal of

bristles on shorter segments; torus greatly enlarged; flagellum strongly plu-

mose; segments 12 and 13 subequal in length; flagellar segment 1 (flavicosta,

flavithorax) or segment 1-4 or 5 with tufts of scales; bristles of flagellar whorls

about 0.4-0.6 length of entire flagellum on proximal segments, about 0.6-1.0

length of segment 13 on that segment. Legs: anterior fore- and midclaws with

basal tooth too small to be seen on pinned specimens, posterior fore- and mid-

claws simple.

Zavortink: Genus Orthopodomyia 93

MALE GENITALIA. Varied, see description for each included group.

PUPAE. Known only for Albipes group.

LARVAE. Varied, consult description for included groups.

BIONOMICS. So far as is known, the immature stages are normally found

in treeholes, bamboo internodes and artificial containers. Females of 2 spe-

cies, albipes and andamanensis, are known to bite man.

SYSTEMATICS. The Orthopodomyia section contains all the Old World spe-

cies except pulchripalpis. These species fall into 1 Oriental and 3 Ethiopian

groups. The Ethiopian species are very poorly known and it is possible that at

least some of them should be placed in the Thomasina section rather than in the

Orthopodomyia section. These possibilities are discussed more fully later (see

especially arboricollis).

It is unfortunate that the isolated species and small groups of Orthopodomy-

ia, which may represent relictual or primitive forms, the study of which is

paramount to understanding the relationships within the genus, are the most

poorly known of all Orthopodomyia. Until the immature stages of the 3 Ethio-

. pian groups and the South American sampaioi are better nom; any classifica-

tion of the genus is provisional.

DISTRIBUTION. Cameroon, India, and southern Japan, south to South Af-

rica and northeastern Australia.

KEYS TO GROUPS

See keys on p. 19-22.

VERNONI GROUP

FEMALES. Head: integument of head capsule, clypeus, torus and flagel-

lum dark brown; eyes separated above antennae by silvery-white to white de-

-cumbent scales; frontal bristles brown; orbital bristles brown, usually 2 later-

al pairs and 3 mesal pairs; lateral and ventral scales white or dingy white; la-

bium considerably swollen apically. Thorax: integument of mesonotum and

postnotum dark brown, scutellum lighter, pleuron evenly dark brown or with

lighter areas; paratergite bare; posterior dorsocentral bristles apparently not

in double row; anterior mesonotal bristles dark brown, supraalar, parascutel-

lar and scutellar bristles lighter; acrostichal scales not separated laterally

from dorsocentral scales by a narrow longitudinal area; pra bristles 1, brown;

upper stp bristles 1, strongly developed, brown, posterior stp bristles 2-4,

amber, weakly to strongly developed, lower stp bristles 5-8, amber, weakly

to strongly developed; upper mep bristles 7-17, whitish; ppn with extensive

area of narrow curved scales; hypostigial scales absent; upper portion of stp

with bare areas at base of bristles, lower stp scales apparently not joined to

those in upper portion; mep with more or less quadrangular patch of flat scales

near margin of stp in upper portion of mep. Legs: integument of forecoxa light,

mid- and hindcoxae brown; forecoxal bristles yellowish-amber, midcoxal bris-

tles brown, hindcoxal bristles brownish-amber; forecoxa with broad flat white

scales on anterior surface and above bristles and with semierect white scales

among bristles, mid- and hindcoxae with broad flat scales in an upper outer

patch and along outer anterior surface; hindtarsal segment 5 entirely dark-

scaled. Wing: scales symmetrical, moderately broad, spreading and clasping

94 | Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

veins; wing largely brown- or black-scaled with white scales forming patches

and scattered, as follows: (1) costal vein with prehumeral, humeral, accessory

sectoral, subcostal and apical light patches and with few to numerous scattered

light scales distally, (2) vein Sc with accessory sectoral and subcostal light

patches, (3) vein R with humeral light patch, sometimes with prehumeral light

patch, and with scattered light scales, (4) vein Rj with accessory sectoral and

apical light patches and with numerous scattered light scales, (5) veins Rg, R4+5,

Cu and 1A with basal light patch, (6) vein R2+3 with light patch opposite base of

vein R4+5, (7) veins M and Cu with light patches distad of crossveins, (8) veins

R92+3 and M with light patch at furcation, (9) all veins behind vein R1 with api-

cal light patch, (10) stem of vein M and veins Cu and 1A with light scales scat-

tered or in small patches, the light scales concentrated in central portion of

stem of M, basad of furcation in Cu, central portion of Cu, and central portion

of 1A. Haltere: integument white to beige, scales transparent to whitish. Ab--

domen: integument of tergites and sternites light brown to brown except for

lighter areas beneath light scales; laterotergite bare; tergite I with largely

dark brown- or black-scaled fan and with a few scatlored light scales. |

MALES. Similar to females. Palpus: 0.96-1.00(?) of proboscis; segment

3 usually with a few strong apical bristles, largely dark brown- to black-scaled,

white scales present at joints between segments 1-2, 2-3 and 3-4 and covering

all of segment 5. Antenna: flagellar segments 1-4 with tufts of extremely long

white or silvery scales.

MALE GENITALIA. Segment VIII: tergite lobe distinct, longer than broad,

apex emarginate and with irregular teeth. Segment IX: tergite with 2 or 3 fine

bristles on each side. Sidepiece: long conical; basal mesal lobe with 4 or 5

stout terete apical bristles which are slightly curved mesally at apex and 1 or 2

finer terete apical bristles sternad of stout ones; mesal surface completely

sclerotized; mesal surface without specialized bristles; tergomesal surface

with strong specialized bristles. Spiniform: normally 1, apex with 3 or 4 ir-

regular lobes. Aedeagus: strongly sclerotized; shape varied; apex with nu-

merous small teeth; each half of sternal surface with large, strong, sharp,

curved spine on distal portion of mesal margin. Proctiger: paraproct sclero-

tization with 2-5 apical teeth; cercal setae 1-3.

PUPAE. Unknown. ;

LARVAE. The one specimen available for study has been provisionally

identified as milloti and is described under that species. See below also.

BIONOMICS. Larvae have been collected in treeholes and bamboo inter-

nodes. Nothing is known of the habits of the adults.

SYSTEMATICS. The Vernoni group contains all the Madagascan species.

To date 3 names, vernoni, milloti and geberti, have been proposed. Accord-

ing to Grjebine (1966:29, 30,32), 7 species of this group occur on Madagascar

and 1 additional species is endemic to Mayote.

The taxonomy of the group is confused. Two species, -vernoni and milloti,

were described as adults and the third species, geberti, was described as a

larva. Van Someren (1949:6-7) described a larva tentatively associated with

vernoni; during this study it was possible to make only a cursory examination

of this larva. I have provisionally associated the only larva available for crit-

ical study with milloti. Since the description of geberti and the description and

my notes of the larva associated with vernoni are quite incomplete, it is not

possible to state with any certainty whether the larvae represent distinct spe-

cies or if they differ from the one larva I have studied. It may be that geberti

is conspecific with the larva associated with vernoni and that the larva illustra-

Zavortink: Genus Orthopodomyia | 995

ted as vernoni by Doucet (1950:57) is conspecific with the one I am associating

with milloti. If this is the case and the tentative associations are correct, then

the larvae of milloti and vernoni might differ on the basis of the length and cur-

vature of the antennae and the length of hair 5-C. However, since both these

features are known to be extremely variable in the Mauritian species arbori-

collis, they may be poor taxonomic characters in this apparently related group.

I have examined adults of vernoni and milloti and find them to be distinct spe-

cies. It appears that the most reasonable procedure to follow here is to recog-

nize these 2 species and to consider geberti as a nomen dubium. The total num-

ber of species involved and the identity of the larvae cannot be ascertained un-

til adults with associated skins are available for study.

Arboricollis appears to be related to this group. Adults of arboricollis and

the Vernoni group agree in wing ornamentation and in the absence of lateroter-

-gite scales and a bare area between the acrostichal and dorsocentral scales.

Larvae of arboricollis and milloti have 6 pairs of hairs in the ventral brush and

stellate body hairs. It is possible, however, that the latter characteristic is

due to parallelism. The other affinities of the Vernoni group are obscure. The

spiniform of the male genitalia is like that of the Neotropical Thomasina section

and the Holarctic Signifera group and the dentition of the aedeagus is reminis-

cent of the Neotropical albicosta. As discussed more fully under arboricollis,

certain features of both arboricollis and the Vernoni group are also found in the

Thomasina section, but it is uncertain whether these similarities indicate re-

lationship or are only retentions of primitive characteristics in separate phyl-

lads. Until the immature stages, particularly the pupa, are better known for

arboricollis, the species in the Vernoni group, and the African Orthopodomyia,

it will not be possible to accurately classify the Ethiopian species, and until

this information is available all the Ethiopian groups are tentatively being as-

sociated with the Albipes group in the Orthopodomyia section.

DISTRIBUTION. Madagascar.

KEYS TO SPECIES

Adults

1. Hindtarsal segment 2 dark-scaled at apex, segment 3 white-scaled in basal

0.5; acrostichal scales mixed brown, white and yellow; dorsocentral

scales largely yellow... . coe oe. TA vernen

Hindtarsal segment 2 with white- scaled aoienl ae ‘seonieut 3 enitely

white-scaled; acrostichal and dorsocentral scales largely white or

Cream *COlOred 2. 6 ae er ee ee ee

Male Genitalia

1. Apex of aedeagus with broad, more or less truncate, lobe on each half;

apical lobe of aedeagus with blunt teeth in more or less distinct rows :

broadest portion of aedeagus parallel sided . ..... .. 11. vernoni

Apex of aedeagus with acute, more or less triangular, lobe on each half;

apical lobe of aedeagus with acute, curved teeth not arranged into dis-

tinct rows; aedeagus-not parallel.sided 2...) se... 4/12. milloti

96 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

11. Orthopodomyia vernoni van Someren

Fig. 18

1949. Orthopodomyia vernoni van Someren, 1949:5-7. *TYPE: holotype o with

genitalia slide 660610-3, Sakaramy, near Diego-Suarez, Madagascar, 1

Apr 1944 [BMNH].

Orthopodomyia vernoni of Knight and Mattingly (1950:2); Hopkins (1952:83-84);

Stone, Knight and Starcke (1959:124); Rickenbach and Hamon (1967:1117).

FEMALE. Unknown.

MALE. Head: decumbent scales silvery anteriorly, yellow posteriorly;

erect scales brown; dorsolateral scales brown anteriorly, white posteriorly;

labium largely brown-scaled, white scales in narrow ring just distad of distal —

end of palpal segment 2, in narrow ventrally incomplete apical ring and in line

from base to middle ring along edge of groove; palpus equal to length of pro-

boscis, segment 3 with scattered white scales. Thorax: acrostichal scales

large, yellow and white at anterior promontory, Small and brown at caudal end,

remainder small, mixed yellow, white and brown; anterior prescutellar scales

small, golden yellow; dorsocentral scales large and white at anterior promon-

tory and at caudal end, remainder small, largely yellow; lateral prescutellar

scales large, brown anteriorly, white posteriorly and laterally; lateral prescu-

tal scales yellow and white; posterior fossal scales largely white; fossal scales

brown; supraalar scales small mesally, larger laterally, yellow and white an--

teriorly, brown posteriorly, silvery-white in middle extension; scutellar scales

white; upper apn scales yellowish; lower apn scales white; ppn scales yellow

above, white below; pra, ppl, pst, ssp, stp and mep scales all white. Legs:

mid- and hindcoxal scales white above, white and brown below; remaining leg

segments with dark scales dark brown or black; femora with dark-scaled pre-

apical ring; forefemur largely dark-scaled, white scales in basal ring and scat-

tered; midfemur anterior surface with largely white-scaled patch basad of dark-

scaled preapical ring, white scales at base ventrally, remainder dark-scaled

basally and ventrally, dark-scaled with white speckles dorsally, posterior sur-

face white-scaled basally, dark-scaled ventrally, dark-scaled with white spec-

kles dorsally, remainder largely white-scaled; hindfemur anterior surface

white-scaled at base ventrally, remainder largely dark-scaled ventrally, dark-

scaled with white speckles dorsally, posterior surface largely white-scaled in

basal portion, largely dark-scaled apically; tibiae with irregular dark-scaled

preapical ring; foretibia with dark-scaled basal ring, anterior, dorsal and ven-

tral surfaces dark-scaled with white speckles, posterior surface largely light-

scaled; midtibia with basal white scales and subbasal dark-scaled ring, ante-

rior, dorsal and ventral surfaces dark-scaled with white speckles, posterior

surface largely white-scaled; hindtibia with narrow white-scaled basal ring and

dark-scaled subbasal ring, dorsal surface dark-scaled with white speckles, an-

terior and ventral surfaces light-scaled with dark speckles, posterior surface

white-scaled in basal portion, remainder dark-scaled; tarsi largely dark-scaled;

foretarsus with white scales in narrow basal ring on segment 1; midtarsus with

white scales in narrow ring at base of segments 1-3; hindtarsus with white

scales in narrow ring at base of segment 1, in moderately broad ring at base

of segment 2, in basal 0.5 of segment 3 and in basal 0.6 of segment 4.. Wing:

vein Ro longer than vein R9,3; vein Mj1,9 longer than M distad of crossvein

m-cu; vein Sc with scattered light scales; vein R415 with light patch in central

Zavortink: Genus Orthopodomyia 97

portion. Abdomen: tergites II-VII largely dark brown- to black-scaled, white

scales in small basolateral patch and in median dorsal or subdorsal patch on

segments II, III, V and VI; tergite VIII with white scales in basal band and apico-~

lateral patch, remainder dark-scaled; sternite II largely bare, white scales

present laterally; sternite II with very large basolateral white patch, remainder

dark-scaled; sternites IV, V largely dark-scaled, white scales in basolateral

patch; sternites VI, VII apparently entirely dark-scaled; sternite VIII largely

dark-scaled, white scales present basally; sidepiece with light and dark scales.

MALE GENITALIA (fig.18). Sidepiece: tergomesal surface with 3 large

stout bristles near basal mesal lobe. Aedeagus: broad, parallel-sided in dis-

tal portion, base narrowed, apex with lateral shoulder, the mesal apical portion

with numerous small blunt teeth arranged into 3 more or less distinct rows.

LARVA. Head: hairs 4-7-C large and with many plumose branches; 13-C

13b, long. Antenna: about 0.5 length of head, straight, swollen in basal 0.5;

Nae be A 3-5b. Thorax: stellate setae present; hair 1-P double; 1-M 8b, Lone:

1-T 5b. Abdomen: stellate setae present; hairs 1, 5,7, 13-II-VI with fewer

branches than in milloti. Siphon: index about 5; hair 1-§ at about middle, 10b.

Anal Segment: hair 1-X double, long; anal gills short.

BIONOMICS. The larva provisionally associated with vernoni was found

in a treehole.

SYSTEMATICS. Some details of the sternopleuron and mesepimeron of

the type of vernoni have been obscured by the minuten pin. One clasper of the

male genitalia from this specimen has 2 spiniforms whereas the other has only

1; I assume that each clasper normally has only 1 spiniform.

The larval description has been taken from van Someren (1949:6-7) and

from notes made at the British Museum of Natural History.

A more complete discussion of the systematics of this species can be found

under the Vernoni group. ;

DISTRIBUTION. Known definitely only from the type locality in extreme

northern Madagascar. Material Examined: 3 specimens; 1, 2 L.

MADAGASCAR. Diego-Suarez: Sakaramy, near Diego- Suarez, 1 Apr 1944,

o holotype [BMNH]; Dec 1944, 2 L [BMNH].

Additional records from the literature. None; see systematics sections

for the Vernoni group and geberti.

12. Orthopodomyia milloti Doucet

Figs. 18, 21

1951. Orthopodomyia milloti Doucet, 1951:66-68. TYPE: holotype %, Périnet,

Tananarive, Madagascar, adult found in a bamboo internode lying on damp

leaves, 1-8 Mar 1950 [IRSM, T.A. 3].

Orthopodomyia milloti of Stone, Knight and Starcke (1959:123); Rickenbach and

Hamon (1967:1117).

Orthopodomyia vernoni of Doucet (1950: 97,62; 1951:64, 65).

FEMALE. Wing: 4.10 mm. Proboscis: 2.51 mm. Forefemur: 2.31 mm.

Abdomen: about 3.0 mm. Head: decumbent scales entirely white or white

mesally and yellow-tinged laterally; erect scales brown laterally and pos-

teriorly, dingy tan near eyes, the remainder dingy cream-colored and

yellow; dorsolateral scales brown anteriorly and white with a tinge of

tan posteriorly; labial-scaling not completely known, dark brown scales

98 | Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

present along dorsal surface in basal 0.4, white scales present dorsally beyond

middle and at apex; palpus straight, about 0.37 of proboscis, 4-segmented,

segment 4 minute, scaling incompletely known, apparently largely dark brown,

with white scales at least at base of segment 3 and with silvery-white scales

covering segment 4; torus scales cream-colored; ilagellar scales medium-

sized, cream-colored. Thorax: acrostichal scales large and white or cream-

colored at anterior promontory, small and brown at caudal end, remainder

small, white or cream-colored; anterior prescutellar scales small, white to

cream-colored or yellowish; dorsocentral scales large and white or cream-

colored at anterior promontory, large and silvery-white at caudal end, remain-

der small, white to cream-colored; lateral prescutellar scales large, brown

anteriorly, silvery-white posteriorly; lateral prescutal and posterior fossal

scales white to cream-colored; fossal scales brown or brown and yellow; su-

praalar scales small mesally, larger laterally, white and cream-colored ante-

riorly, brown posteriorly, silvery-white in middle extension; scutellar scales

_ silvery-white; apn, ppn, ppl, pst, ssp, stp and mep scales white to cream-col-

ored; pra scales white to dingy white. Legs: mid- and hindcoxal scales white

or brown and white; remaining leg segments with dark scales dark brown to

black; femora with dark-scaled preapical ring; forefemur largely dark-scaled,

white or cream-colored scales in basal ring, on distal portion of posterior sur-

face and scattered; midfemur anterior surface with cream-colored patch basad

of preapical dark-scaled ring, white scales at base ventrally, remainder dark-

scaled basally and ventrally, dark-scaled with cream speckles dorsally and api-

cally, posterior surface white-scaled basally, dark-scaled dorsally and in basal

ventral portion, remainder largely pale-scaled; hindfemur anterior surface

white-scaled at base ventrally, remainder dark-scaled ventrally, dark-scaled

with cream speckles dorsally, posterior surface white-scaled at base, dark-

scaled ventrally and apically, dark-scaled with cream speckles dorsally, re-

mainder largely pale-scaled; foretibia anterior and dorsal surfaces dark-scaled

with cream-colored speckles, posterior and ventral surfaces largely pale-

scaled; midtibia with mostly cream-colored scales basally except for dark-

scaled subbasal ring, remainder dark-scaled with cream-colored speckles be-

coming entirely dark-scaled preapically; hindtibia with mostly cream-colored

scales basally except for irregular dark-scaled ventrally incomplete subbasal

ring, dark-scaled preapically, remainder largely dark-scaled with cream-col-

ored speckles on anterior and dorsal surfaces, largely light-scaled on posteri-

or and ventral surfaces; tarsi largely dark-scaled; foretarsus with white scales

in small dorsal patch or complete ring at base of segment 1, in small dorso-

basal patch on segment 2 and sometimes on segment 3; midtarsus with white

scales at base of segments 1-4, those on segments 1, 2 and sometimes 3 form-

ing a complete ring, those on segment 4 in dorsal patch; hindtarsus with white

scales in small dorsobasal patch on segment 1, in narrowbasal and broader api-

cal ring on segment 2, covering all of segment 3 and in basal 0.67 of segment 4.

Wing: vein Ro 1.67 length of vein Ro.3; vein M149 1.14 length of M distad of

crossvein m-cu; vein Sc apparently without scattered light scales; vein R4,5

apparently without light patch in central portion. Abdomen: tergites H-VIII

largely dark brown-scaled with white scales in basolateral patch and possibly

in dorsobasal or median subdorsal patch; sternite II largely white-scaled,

brown scales present apically and mesally; sternites II-VII largely brown-

scaled, with white scales in basolateral patch; sternite VIII largely bare, white

scales present basolaterally.

MALE. Essentially as infemale. Proboscis: largely dark brown- or

Zavortink: Genus Orthopodomyia | 99

black-scaled, white scales in narrow ring just distad of distal end of palpal seg-

ment 2, cream-colored scales at apex laterally. Palpus: about 0.96 of probos-

cis, apparently without scattered light scales on segment 3. Antenna: integu-

ment of torus lighter latérally. Abdomen: light-scaling possibly more extensive

than in female; tergite VIII with narrow basal light band; sternite VIII dark-

scaled with basolateral and apicolateral light patches; sidepiece with light and

dark scales.

MALE GENITALIA (fig. 18). Sidepiece: tergomesal surface with 1 stout

and 2 finer specialized bristles near basal mesal lobe. Aedeagus: middle por-

tion broad, with sides diverging distally, base narrow, apex without lateral

shoulder, but with each half narrowing to acute tooth-bearing lobe, teeth nu-

merous, curved and sharp-pointed, not arranged into rows.

LARVA (fig.21). Head: 1.25 mm. Siphon: 1.81 mm. Anal Saddle: 0.54

mm. Head: broader than long; integument not spiculose; largely tan colored,

lighter around imaginal eye, darker posteriorly, collar brown; labrum short;

mental plate brown, with 10,11 teeth on each side; hair 2-C apparently absent;

4-C long, extending beyond apex of antenna; 13-C 6, 7b, long; 14-C moderately

developed, 2,3b; maxillary sclerite hair strongly developed, 3,4b. Antenna:

moderately-long, narrow, curved mesally; integument not spiculose; uniformly

brown colored; hair 1-A 7,8b. Thorax: epidermal pigment strongly developed,

purple in color; spicules not conspicuous; hairs strongly pigmented; hairs 0-P,

1,14-M and 1,3,4-T stellate; 1-P multiple; 1-M,T 12-14b; 4-M single. Abdo-

men: segments VI, VII without sclerotized plates; hairs strongly pigmented;

hair 1-I, II stellate; 2-I, II laterad of hair 1 of corresponding segment; 6-I, II

multiple, 6-III-VI single; 12,13-I stellate, branches of 13-I shorter than 10-I;

13-II stellate; 1-III, IV, VI, VII moderately long, multiple; 5-III-VI moderately

long, multiple; 13-III, IV, VI, VII moderately long, multiple; 1-V very long, 3b;

13-V long, multiple; 2-VI, VII laterad of hair 1 of corresponding segment; 8-

VII stellate. Segment VII: sclerotized plate present; comb scales in 2 rows;

anterior comb scales about 26, posterior comb scales about 15; larger anterior

comb scales shaped like posterior comb scales, with 1 long apical spine and

basal fringe; hairs 1, 5-VIII strongly developed, stellate. Siphon: index about

4.8; brown, with extreme base darker, apex lighter, portion distad of alveolus

of hair 1-S slightly darker than portion basad of 1-S; integumentary sculpturing

moderately developed basally, imbricate; hair 1-S located about 0.44 from base

of siphon, 11b, longest branches reaching about 0.4 distance from alveolus to

apex of siphon. Anal Segment: saddle complete; sclerotized incomplete ring

present basad of saddle; integumentary sculpturing weakly developed, imbri-

cate; brown, extreme base darker, dorsal portion slightly darker than ventral

portion; hair 1-X 2,3b, long; ventral brush apparently with 6 pairs of hairs;

gills short, dorsal pair 1.5 times length of ventral pair and 0. 85 length of anal

saddle. |

BIONOMICS. Adults of the type series were collected in bamboo internodes

lying on the ground. Larvae have also been collected in bamboo internodes.

SYSTEMATICS. Milloti is distinct from vernoni on the basis of the adult

and male genitalic characters given in the keys.

The small sample of adults available for examination is quite uniform, with

only some variation in the sizes of the light patches and the number of scattered

light scales on the wing. The lateral prescutal, posterior fossal and anterior

prescutellar scales are cream-colored to yellowish and apparently not as strong-

ly yellow-pigmented as in the type series.

The larva described here was collected at the type locality of milloti and I

100 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

am provisionally associating it with this species. It agrees well with the larva

identified as vernoni and figured by Doucet (1950:57). It differs from the larva

described by geberti and figured by Grjebine (1953:465-467) by having longer,

curved antennae and longer hair 5-C.

DISTRIBUTION. Known definitely only from east-central Madagascar.

Material Examined: 6 specimens; 4c, 19, 1L.

MADAGASCAR. Tamatave: Fenerive, 24 Dec alata Sakondroely, 40°, 192

[TERT].

Tananarive: Périnet, 1 L [USNM].

Additional records from the literature. MADAGASCAR. Diego-Suarez:

Montagne d'Ambre, R. Paulian (Doucet, 1950:57,62). Larvae collected at this

locality were reported as vernoni, but may be milloti; see RySTEMeHICS discus-

sion of milloti.

NOMEN DUBIUM

12a. Orthopodomyia geberti Grjebine

1953. Orthopodomyia geberti Grjebine, 1953:465-467. TYPE: holotype larva,

forest station at Ampijoroa, Ambato-Boeni District, [Majunga], Mada-

gascar, larva found in a treehole, 10 Dec 1951 [IRSM].

Orthopodomyia geberti of Stone, Knight and Starcke (1959:123); Rickenbach and

Hamon (1967:1117).

SYSTEMATICS. Grjebine described geberti on the basis of the larva alone.

He apparently believed it differed from the larva described and tentatively as-

sociated with vernoni by van Someren (1949:6-7) because it did not have stellate

hairs on the thorax and body. This difference may well be a matter of Seman-

tics, since geberti was stated to have ''tufts of strong plumose hairs.'' Com-

parison of the descriptions of Grjebine and van Someren fails to show any means

of distinguishing the larvae and raises the possibility that geberti is a synonym

of vernoni. However, since the larva described by van Someren was only pro-

visionally considered to be vernoni, and since both descriptions are rather in-

complete, I think it better to consider geberti as a nomen dubium than a syno-

nym.

DISTRIBUTION. Type from Ampijoroa, Majunga, Madagascar. Specimens

Examined: none.

NKOLBISSONENSIS GROUP

13. Orthopodomyia nkolbissonensis Rickenbach and Hamon

1967. Orthopodomyia nkolbissonensis Rickenbach and Hamon, 1967:1112-1117.

TYPE: holotype 2, Center for Agricultural Research of Nkolbisson, 8 km

from Yaoundé, south Cameroon, adult found in lower vegetation of forest,

18 Jan 1965 [IERT].

FEMALE. Head: integument of torus yellow; decumbent scales white to

pure yellow; erect scales pale mesally, darker laterally; labium largely black-

scaled, pale yellow scales in a broad ring distad of middle and at apex (?);

Zavortink: Genus Orthopodomyia 101

palpus almost 0.95 of proboscis, largely black-scaled, with pale yellow scales

in median ring and narrow zone at distal end; torus and flagellar scales pale.

Thorax: mesonotum deep-.brown, almost black, pleuron dark; paratergite bare;

acrostichal scales separated from dorsocentral scales by a longitudinal bare

area; scales bright yellow to middle of scutum where dense black scales (ante-

rior lateral prescutellar scales, ?) form a pair of large triangular patches lat-

erad of dorsocentral bristles; scutellar scales bright yellow; apn and ppn scales

_ yellow; ppl, stp and mep scales cream-colored; mep with middle patch of

scales. Legs: coxae with patch of cream-colored scales on upper part; femora

with yellow scales at apex; forefemur with black scales at base, in preapical

ring, and scattered in basal 0.67 of anterior surface, remainder yellow-scaled;

midfemur with black scales at base, in preapical ring, in streak in middle por-

tion of dorsal surface and densely scattered in middle portion of anterior sur-

face, remainder yellow-scaled; hindfemur with black scales at base and in pre-

apical ring, remainder yellow-scaled; tibiae yellow-scaled with black scales in

broad subbasal, median and preapical rings and sometimes on ventral surface

basally; tarsi largely black-scaled; tarsal segment 1 with yellow scales in bas-

al, median and apical rings, the median ring broad on the mid- and hindlegs;

tarsal segment 2 with yellow scales in basal and apical rings; tarsal segment 3

with yellow scales in narrow basal and apical rings, either or both rings some-

times reduced to patches or absent on foreleg; tarsal segments 4 and 5 usually

entirely black-scaled, yellow scales sometimes present in basal zone on seg-

ment 4 or over entire surface of segment 5. Wing: costal vein, veins Sc and R

and bases of veins M, Cu and 1A with large scales, otherwise scales long and

narrow; scales black and cream-colored; costal vein black-scaled in basal

0.67; basal 0.33 of wing with mixed cream-colored and black scales, with either

color predominating, middle 0.33 black-scaled, apical 0.33 with cream-colored

scales only or with some black scales. Haltere: yellow. Abdomen: tergites II-

VII largely black-scaled, cream-colored scales in basolateral oblique patch,

yellow scales connecting patches of opposite sides, forming a basal band convex

towards the base on tergite II and a preapical band convex towards the apex on

tergites V-VII, tergite VII sometimes with light streak from base to preapical

band middorsally; tergite VIII with pale scales basally; sternites largely dark-

scaled, yellow scales forming basal band.

MALE. Similar to female. Proboscis: largely black-scaled, yellow scales

in complete median ring, sometimes in ventrally incomplete ring nearer apex,

and streaked along dorsal surface from distal ring to apex or from median ring

to near apex. Palpus: about as long as proboscis, ornamentation apparently as

follows: segment 1 and possibly base of segment 2 black-scaled with yellow

speckles, segment 2 yellow-scaled, segments 3 and 4 black-scaled except for

yellow scales over joint between them, segment 5 pale yellow-scaled.

MALE GENITALIA. Segment IX: tergite with 4-6 bristles on each side,

these apparently quite strong. Sidepiece: long conical; basal mesal lobe with

4 or 5 stout terete apical bristles which are curved mesally at apex and appar-

ently with some long finer bristles sternad of stout ones; mesal surface possi-

bly with moderately developed specialized bristles; tergomesal surface with ©

strong specialized bristles near basal mesal lobe. Spiniform: 1, simple. ae

deagus: strongly sclerotized, acorn-shaped, apparently without sternal teeth.

Proctiger: paraproct sclerotization with only 1 apical tooth; cercal setae 4.

PUPA, LARVA. Unknown.

BIONOMICS. Adults were captured in the lower vegetation of a forest.

SYSTEMATICS. Nkolbissonensis was the first species to be described

102. - Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

from the continental portion of the Ethiopian Region. It appears to be very dis-

tinct from all other Orthopodomyia on the basis of the male genitalia and adult

ornamentation. The immature stages are unknown.

I have not examined any specimens of nkolbissonensis and the description

provided above has been taken from Rickenbach and Hamon (1967:1112-1117).

This species has been given group rank and placed in the Orthopodomyia

section. The latter action is taken on the basis of geography only.

DISTRIBUTION. Type series from Yaoundé, south Cameroon. Specimens

Examined: none.

14. Orthopodomyia sp. South African form

Orthopodomyia species of Mattingly and Brown (1955:104); Muspratt (1955:156, ©

194). ,

SYSTEMATICS. Muspratt reported finding 1 damaged adult and some lar-

vae of an undescribed species of Orthopodomyia in a treehole at Satara Camp,

Kruger National Park, South Africa. All that is known about this species is the

following statement taken from the key presented by Muspratt: 'medium-sized

blackish species, proboscis with some pale scales, but unbanded, femora and

tibiae speckled and with white bands on the tarsi, some of which embrace the

joints." | : 7

This species cannot be described until the original specimens are located

or additional material is collected. Its relationships are, of course, unknown;

it is tentatively being associated with nkolbissonensis, the only species de-

scribed from mainland Africa. : |

DISTRIBUTION. Known only from Satara Camp, Kruger National Park,

South Africa. Specimens Examined: none. 7

ARBORICOLLIS GROUP

15. Orthopodomyia arboricollis (Charmoy)

o Wiss, 18°22

1908. Culex arboricollis Charmoy, 1908:257-258. *TYPE: lectotype & (148),

specimen labeled as "Type o,'' Vacoas, Mauritius, reared from a larva

found in a treehole, 1908, R. Ross; PRESENT SELECTION [LIVER in

BMNH].

Orthopodomyia arboricollis of Edwards (1920:135); MacGregor and Gebert

(1923:449-452); MacGregor (1927:142-147); Edwards (1932:107, 108); Hop-

kins (1936:62-63); Edwards (1941:72-73, 374); van Someren (1949:5, 6);

Knight and Mattingly (1950:2); Hopkins (1952:82-83); Halcrow (1954:239);

Mattingly and Brown (1955:104); Stone, Knight and Starcke (1959:122); Rick-

enbach and Hamon (1967:1117). |

Newsteadina arboricollis of Theobald (1909:297; 1910b:475-477).

FEMALE. Wing: 3.64 mm. Proboscis: 2.05mm. Forefemur: 2.12 mm.

Abdomen: about 2.5 mm. Head: integument of head capsule, clypeus, torus

Zavortink: Genus Orthopodomyia 103

and flagellum brown to dark brown; eyes separated above antennae by yellowish

to yellow scales; frontal bristles brown; orbital bristles brown, usually 3 lat-

eral pairs and 2 mesal pairs; decumbent scales silvery-white to cream-colored

or yellowish; erect scales behind eyes narrow, brown, those on occiput broad-

er, brown posteriorly and laterally, remainder yellowish; dorsolateral scales

brown anteriorly, cream-colored to yellowish posteriorly; lateral and ventral

scales cream-colored to yellowish; labium considerably swollen apically, large-

ly dark brown- to black-scaled, white or slightly yellow-tinged scales in narrow

ring about 0.6-0.7 from base and scattered; palpus 0.45 of proboscis, 4-seg-

mented, segment 4 short, largely dark brown- to black-scaled, white scales at

joints between segments 1-2 and 2-3, covering all of segment 4 and scattered;

torus scales white or dingy; flagellar scales small and medium-sized, whitish.

Thorax: integument of mesonotum, postnotum and pleuron uniformly brown to

dark brown or posterior portion of mesonotum, postnotum and posterior portion

of pleuron lighter, scutellum always lighter; paratergite bare; posterior dorso-

central bristles apparently not in double row; anterior mesonotal bristles dark

brown, supraalar, parascutellar and scutellar bristles brownish-amber; acros-

tichal scales not separated laterally from dorsocentral scales by a narrow lon-

gitudinal area, large white or cream-colored scales at anterior promontory and

sometimes in interrupted or more or less continuous row throughout length of

scale band, large or small brown scales at posterior end, remaining scales

small, white and yellowish or all yellow; anterior prescutellar scales small,

golden; dorsocentral scales large, white or cream-colored at anterior promon-

tory, large, silvery-white and yellowish at posterior end, remainder small,

silvery-white anteriorly, remainder yellow or yellow and brownish; lateral

_ prescutellar scales large, brown anteriorly, silvery-white posteriorly; lateral

prescutal scales silvery-white anteriorly, remainder largely dingy-yellow, of-

ten with brown or pink reflection; posterior fossal scales dingy-white and din-

gy-yellow, often with pink reflection; fossal scales brown and dingy-yellow,

sometimes with pink reflection; supraalar scales small mesally, larger later-

ally, posterior scales brown, remainder whitish, silvery-white and dingy-yel-

low, often with pink reflection; scutellar scales silvery-white; pra usually with

1 brown bristle (0-2); upper stp bristles 1, strongly developed, brown, poste-

rior stp bristles amber, 2 or 3 strongly developed, 1 or 2 moderately devel-

oped, lower stp bristles 3-7, amber, weakly to strongly developed; upper mep

bristles 7-14, white or rarely yellowish; upper apn scales dingy yellow; lower

apn scales cream-colored with yellow reflection; ppn with almost complete cov-

ering of dingy white and dingy yellow narrow curved scales, scales sometimes

with pink reflection; pra, ppl, pst and ssp scales cream-colored with yellow

reflection; hypostigial scales absent; upper half of stp with bare areas below

pra and at bases of bristles, lower posterior stp scales joined to those in upper

half, all stp scales cream-colored with yellow reflection; mep with flat scales

in a lower anterior triangular patch which may be connected to upper mep scale

patch, scales cream-colored with yellow reflection. Legs: integument of mid-

and hindcoxae same color as pleuron, forecoxa lighter; forecoxal bristles yel-

lowish-amber, midcoxal bristles brown, hindcoxal bristles brownish amber;

coxal scales largely broad, flat, the light ones cream-colored with yellowish

reflection; forecoxa with light scales on anterior surface, above and among

bristles and in small patch on lower posterior surface, some of those among

bristles semierect, midcoxa with patch of light scales above bristles and with

light and brown scales on anterior surface, hindcoxa with scales on anterior

surface, those above light, those below brown or brown and light; remaining

104 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

leg segments with dark scales brown or black, light scales white to cream-col-

ored with yellowish tinge; femora with dark-scaled preapical ring; forefemur

with light-scaled streak at base of dorsal surface, remainder dark-scaled with

scattered light scales; midfemoral anterior surface dark-scaled at base and

ventrally in basal 0.2-0.5, remainder dark-scaled with scattered light scales

which become more numerous apically, posterior surface dark-scaled ventral-

ly to beyond middle, usually with pale basal streak, remainder dark-scaled with

scattered light scales; hindfemur like midfemur except that ventral streak of

dark scales on both anterior and posterior surfaces extends further apically;

tibiae with dark-scaled irregular preapical ring; foretibia with dark-scaled bas-

al ring, anterior and dorsal surfaces dark-scaled with scattered light scales,

posterior surface with light-scaled streak, ventral surface entirely dark-scaled;

midtibia with a few light scales at base followed distally by dark-scaled ring,

dorsal, anterior and ventral surfaces dark-scaled with scattered light scales,

posterior surface with light-scaled streak; hindtibia with narrow irregular bas-

al light-scaled ring followed distally by irregular dark-scaled subbasal ring,

dorsal and anterior surfaces dark-scaled with scattered light scales, ventral

surface largely light-scaled, posterior surface largely dark-scaled or with light

speckles; tarsi largely dark-scaled, segment 1 with incomplete to complete nar-

row basal light-scaled ring; foretarsal segment 1 dark-scaled at apex, seg-

- ments 2-5 all dark-scaled; midtarsal segment 1 with a few light scales at apex,

segments 2-3 or 4 with a few light scales at base; hindtarsal segment 1 with

moderately broad apical light-scaled ring, segments 2 and 3 with moderately

broad basal and narrow apical light-scaled rings, segment 4 with light-scaled

ring in basal 0.5, segment 5 entirely dark-scaled or with a few light scales at

apex. Wing: vein Ro 3.55 length of vein Rog; vein My.9 1.86 length of vein

M distad of crossvein m-cu; scales mostly asymmetrical, moderately to quite

broad, spreading on most veins; wing largely dark brown- to black-scaled, with

light, usually white, scales in definite but variable-sized patches and scattered,

as follows: (1) costal vein with basal, prehumeral, humeral, accessory sector-

al, subcostal and apical light patches and with few to numerous scattered light

scales, (2) vein Sc with accessory sectoral and subcostal light patches and with

few to numerous scattered light scales, (3) vein R with humeral and sometimes

prehumeral light patches, and with few to numerous scattered light scales, (4)

vein R; with accessory sectoral and apical light patches and with few to numer-

ous scattered light scales, (5) veins Rg, R4,5, Cu, 1A and sometimes M with

basal light patch, (6) vein Ro9,3 with light patch opposite base of vein Rq4,5, (7)

veins M and Cu with light patches distad of crossveins, (8) all veins behind vein

Rz with apical light patch, (9) veins Ro, R3, R45, M, Cu and 1A with few to

numerous light scales in small patches or scattered, (10) portion of costa along —

posterior edge of wing with light scales in patches at ends of veins and scattered.

Haltere: integument white to beige, scales transparent to whitish or yellowish.

Abdomen: integument of tergites and sternites light brown to brown except for

lighter areas beneath light scales; laterotergite bare; tergite I with brown or

brown and light scales in fan, with lateral patch of light scales and with scat-

tered light and dark scales; remaining segments with dark scales brown to dark

brown or black; tergites II-VIJ with white scales in large basolateral patch and

usually in narrow irregular basal band, patch on segments II and III sometimes

extending to apex of segment near lateral edge of tergite, patch on each suc-

ceeding segment smaller, basal band of segment II sometimes broadened mid-.

dorsally; tergite VIII with white scales in basal band; sternite II largely bare,

with a few white scales; sternite III largely white-scaled, brown scales mesally

| Zavortink: Genus Orthopodomyia 105

and in narrow apical band; sternites IV-VII with white scales in basolateral patch

and sometimes in irregular narrow basal band, patch smaller on each succeed-

ing segment; sternite VIII bare mesally, brown-scaled laterally.

MALE. Essentially as in female. Head: mesal posterior erect scales yel-

lowish. Proboscis: largely dark brown- or black-scaled, white or yellow-

tinged scales in narrow to broad ring just distad of distal end of palpal segment

2 and scattered. Palpus: about 0.92 of proboscis, segment 3 with a few strong

apical bristles; largely dark brown- or black-scaled, white scales at joints be-

tween segments 1-2, 2-3 and 3-4 and scattered on segments 2 and 3, white or

silvery-white scales sometimes at apex of segment 4 and always covering all

of segment 5. Antenna: integument of torus lighter laterally; flagellar segments

1-4 with tufts of extremely long silvery-white scales. Abdomen: basal light i

bands of tergites better developed than in female; sternite VIII white-scaled

apically and laterally, remainder dark-scaled; sidepiece with light and dark

scales.

MALE GENITALIA (fig. 18). Segment VIII: tergite lobe distinct, slightly

longer than broad, apex truncate or shallowly emarginate, apical portion (not

just margin) with small teeth. Segment IX: tergite with 2 or 3 fine bristles on

each side. Sidepiece: long conical; basal mesal lobe with 4 stout terete apical

bristles which are curved mesally at apex and 2 finer terete apical bristles

sternad of stout ones; mesal surface more or less completely sclerotized; mes-

al surface with moderately developed specialized bristles, bristles long, curv-

ing sternally and basally at apex; tergomesal surface sometimes with 1 or 2

poorly developed specialized bristles distad of basal mesal lobe. Spiniform: 1,

simple. Aedeagus: moderately sclerotized; central portion broadest, tapering

to both a narrow base and apex; apex slightly flared, the edge of the flare wrin-

kled; sternal surface with tooth on each half along mesal edge near apex. Proc-

tiger: paraproct sclerotization with 3 or 4 apical teeth; cercal setae 2-4.

PUPA. Not available for study. According to MacGregor and Gebert

(1923:452) hair 9-VIII is a ''dense plume" and is considerably longer than hair

9-VII.

LARVA (fig. 22). Head: 1.04 mm. Siphon: 0.93 mm. Anal Saddle: 0.38

mm. Head: broader than long, integument not spiculose; light tan to tan in col-

or, lighter around imaginal eye, darker posteriorly, collar brown; labrum

short; mental plate tan, with 8-10 teeth on each side; hair 2-C apparently ab-

sent; 4-7-C apparently extremely variable in size and number of branches;

13-C strongly developed, 15-24b in specimens available for study. Antenna:

apparently short and straight in individuals with hairs 4-7-C reduced, moder-

ately-long and curved in individuals with hairs 4-7-C strongly developed; integ-

ument without spicules; uniformly light tan to tan. Thorax: epidermal pigment

absent in specimens examined; spicules not conspicuous; hairs strongly pig-

mented and strongly plumose; chaetotaxy extremely variable; hairs 0-P, 1, 14-

M and 1,3,4-T stellate; hairs 1,3,4,8-P, 13-M and 8-T somewhat stellate; 5-

P and 6-M single or multiple; 4-M always single. Abdomen: segments VI, VII

with or without sclerotized plates; hairs strongly pigmented and strongly plu-

mose; chaetotaxy extremely variable; hairs 1,5,13-I-VII, 7-II-VI, 8-VI, VII,

9-III-VI and 12-I stellate; 2-I, II not stellate, laterad of hair 1 of corresponding

segment; 6-I multiple, 6-II-VI double; branches of 13-I shorter than 10-I; 2-

III-VII laterad of hair 1 of corresponding segment, moved cephalad and stellate.

Segment VIII: sclerotized plate present; comb scales in 2 rows; anterior comb

scales 26-31, posterior comb scales 9-14; larger anterior comb scales shaped

like posterior comb scales, with 1 long apical spine and basal fringe; hairs

106 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

1,5-VIII stellate; hair 3-VIII only moderately developed, 3-7b. Siphon: index

2.9-3.2; largely tan or brown with darker base and lighter apex; integument

ornamented with spinules which are more conspicuous basally; hair 1-S located

about 0.33-0.38 distance from base, only moderately developed, 3-5b, longest

branches not reaching apex of siphon. Anal Segment: saddle complete; sclero-

tized incomplete ring at base of saddle present or absent; integument with spin-

ules; tan to brown, darker dorsally and at base; hair 1-X strongly developed,

multiple; ventral brush with 6 pairs of hairs, the more caudal hairs with 8-12

branches; gills very short, rounded, dorsal pair about 1.2 times length of ven-

tral pair and about 0.45 length of anal saddle.

BIONOMICS. The immature stages are found in treeholes. MacGregor and

Gebert (1923:450) report that the larvae are very common. Nothing is known of

the habits of the adults. MacGregor (1927:142) states that except for specimens

which have just emerged from the pupa, adults have never been encountered in

nature.

SYSTEMATICS. This is the most beautifully ornamented species of Ortho-

podomyia and it is one of the most interesting from the aeepoate of evolution

and zoogeography.

The adults of arboricollis are distinctive on the basis of mesonotal, wing

and hindtarsal ornamentation. They are rather uniform, with conspicuous var-

iations involving only the mesonotal and wing ornamentation. On the mesonotum

the acrostichal scales range in color from white to deep yellow and the propor-

tions of the different colors of scales making up the lateral prescutal, fossal, .

posterior fossal and supraalar scale areas differ. The wing exhibits variation

in the size of the light patches and in the number of scattered light scales.

The larva of arboricollis is one of the most characteristic in the genus.

The form of hairs 6-II-VI and the possession of spinules on the siphon and anal

saddle are unique. The remainder of the larval chaetotaxy is also quite distinct.

Some of the hairs, such as the relatively poorly developed 3-VIII and 1-S, are

apparently neotenic retentions while others are strongly developed and stellate.

In spite of the limited distribution of this species and the uniformity of the adults,

the larva of arboricollis is evidently the most variable in the genus. In the lim-

ited number of specimens available for study it appears that larvae from any

one collection are quite uniform in chaetotaxy but may differ considerably from

those of another collection. A particular hair, for example 6-M, may be single

in individuals in one collection, 3-5 branched in those from another treehole

and 9-11 branched in yet another series of larvae. The number of branches in

the stellate hairs varies widely and again individuals from any particular col-

lection display only a limited portion of the variation. These observations indi-

cate that hairy forms are environmental modifications.

I have not seen larvae with either as hairy or non-hairy head capsules as

those studied by MacGregor and Gebert (1923:451). These authors illustrate a

specimen with short, straight antennae and hairs 4-7-C weakly developed and

another with long, curved antennae and hairs 4-7-C strongly developed. These

authors indicate that both the hairy and non-hairy larvae they illustrated came

from a single treehole and that intermediates between the 2 extremes existed

in the same collection. This would imply that there is a genetic component in-

volved in determining the hairiness of an individual or that the treehole environ-

ment may be heterogeneous in time and conditions existing during some interval

stimulate hairiness in only those individuals that pass through a particular stage

in their development during that interval.

Although many other species of Orthopodomyia have non-hairy and hairy

Zavortink: Genus Orthopodomyia — 107

larval forms, arboricollis is the only one in which even the head hairs are in-

volved. The "non-hairiest" larva available for study still had most body hairs

stellate; it is not known if this would be the case in a larva with reduced head

hairs.

Edwards (1932:108) placed arboricollis in Group A (Orthopodomyia) along

with the Oriental species, the Central American phyllozoa and the Neotropical

fascipes. When Knight and Mattingly (1950:1-2) subdivided Edward's Group A,

they erected the arboricollis subgroup for arboricollis, the Madagascan verno-

ni and the Indian species flavicosta and flavithorax on the basis of the long male

palpi and the absence of an accessory subcostal light patch on the wing. A more

detailed study of the arboricollis subgroup of these authors has shown that it is

an unnatural assemblage. Even on adult characters flavicosta and flavithorax

are more similar to the species in the anopheloides subgroup of Knight and

Mattingly than to arboricollis or vernoni and a study of the immature stages has

shown that at least one of them, flavicosta, is definitely allied with the anophe-

loides subgroup of these authors.

An appraisal of the affinities of arboricollis is difficult because of the mea-

ger information available on the immature stages of it and the Ethiopian species.

Adults of arboricollis have the mesonotum nearly completely covered with scales

and in this respect they are similar to adults in the Neotropical Thomasina sec-

tion, the Madagascan Vernoni and Oriental Albipes groups, and the African nkol-

bissonensis. A closer relationship between arboricollis and the species in the

Thomasina section and the Vernoni group is suggested by the fact that all of

these species do not have a longitudinal bare area separating the acrostichal and

dorsocentral scale bands and all have a single upper stp bristle. In addition,

arboricollis and the species in the Thomasina section are the only species of

Orthopodomyia with very broad asymmetrical wing scales, and arboricollis and

_ the species in the Vernoni group are the only ones among those species with an

extensively scaked mesonotum lacking scales on the laterotergite and possessing

an accessory Sectoral light patch on the costa.

The male genitalia of arboricollis are very Similar to some species in the

Albipes group. They are dissimilar to both the Vernoni group and .the Thomas-

ina section.

As indicated before, the pupa is the most reliable stage for showing affini-

ties in Orthopodomyia, and until it is adequately described for this species all

hypotheses on the relationships of arboricollis must be provisional.

The larva of arboricollis is superficially similar to vernoni in possessing

numerous stellate body hairs. While this may indicate true relationship, since

this type of chaetotaxy has been evolved separately in larvae of many different

genera of mosquitoes, one must consider that these similarities may only be

due to parallelism. Two characteristics of the chaetotaxy of Orthopodomyia

larvae have been found to be of particular value in hypothesizing group relation-

ships. The first of these is the form of hair 4-M. This hair is always single

in Species in the Holarctic Signifera group and the Thomasina section and it is

normally always double or multiple in the Albipes group and the Central Amer-

ican phyllozoa. The second characteristic is the position of hairs 2-VI, VII. In

the Signifera group and in at least fascipes of the Thomasina section these hairs

are laterad of hair 1 of the corresponding segment, whereas in the Albipes

group and phyllozoa they are mesad of hair 1. In arboricollis and milloti hair

4-M is single; this again suggests a closer relationship to the Thomasina sec-

tion than to the Albipes group. The position of hairs 2-VI, VII cannot be used to

_ indicate affinities in arboricollis because in association with their stellate de-

108 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

velopment these hairs have moved far cephalad and laterad of hair 1 on all of

abdominal segments III-VII. In milloti hair 2 is laterad of hair 1.

Arboricollis appears to be distinctive enough to be placed in a monotypic

group. I believe its closest affinities are with the Vernoni group, and for the

time being I am placing arboricollis, the Vernoni group and nkolbissonensis in

the Orthopodomyia section largely on the basis of geography. This arrange-

ment is provisional only and is subject to change when the pupae of arboricollis

and the Vernoni group and the immature stages of the continental African spe-

cies are described.

As indicated above, some features of both arboricollis and the Vernoni

group are found in the Neotropical Thomasina section. It is possible that these

similarities are not indicative of relationship but are retentions of primitive

characteristics by distinct groups. There is some evidence that at least 1 of

the characteristics shared by the 3 groups, the presence of an upper stp bris-

tle, is primitive and consequently its presence cannot be used to unite these

eroups.

DISTRIBUTION. Endemic to Mauritius. Material Examined: 33 specimens;

Wop Geo i |

MAURITIUS. Réduit, M.E. MacGregor, 4 July 1922, 1 L [BMNH]. Va-

caos, 1908, R. Ross, & lectotype (148), 2? [LIVER in BMNH]. No locality,

15 Sept 1922, M. E. MacGregor, 2 & [BMNH]; Sept-Nov 1922, M.E. MacGre-

gor, 1o, 19 [BMNH]; Mar-June 1952, 1%, 59, 5L [IERT]; Jan 1953, 4 L

[IERT]; 27 Sept, M.E. MacGregor, 2 L [BMNH]; 1952, 20°, 12 [UCLA]. No

locality or date, M. E. MacGregor, 22, 3 L [BMNH]. | 3

Additional records from the literature. None.

ALBIPES GROUP.

FEMALES. Head: eyes separated above antennae by white to cream-col-

ored scales; orbital bristles usually 1 or 2 lateral pairs and 2-4 mesal pairs;

labium slightly swollen apically; palpus 0.42-0.58 of proboscis, 4- or 5-seg-

mented; flagellar scales large and small. Thorax: paratergite with (flavicos-

ta, flavithorax) or without scales; posterior dorsocentral bristles sometimes

ina double row; anterior mesonotal bristles brownish-amber to dark brown,

supraalar, parascutellar and scutellar bristles lighter, yellowish-amber to tan;

acrostichal scales separated from dorsocentral scales by a longitudinal bare

area, large and white or cream-colored at anterior promontory, remainder

small; anterior prescutellar scales usually small, sometimes a few large and

elongate, all golden or yellow or with a few brown posteriorly; dorsocentral

scales large and white or cream-colored at anterior promontory, large at cau-

dal end, remainder small; lateral prescutellar scales large, brown anteriorly;

lateral prescutal scales white or rarely a few yellowish; supraalar scales usu- |

ally small in anterior mesal area, remainder larger; scutellar scales white to

silvery-white; pra bristles absent or 1, brown; upper stp bristles absent, pos-

terior stp bristles 1-3, amber, moderately to strongly developed, lower stp

bristles 0-7, amber, weakly to moderately developed; upper posterior mep

bristles 3-10, yellowish-amber (flavicosta, flavithorax) or white, moderately

developed, when white they may be somewhat flattened and difficult to distin-

gcuish from scales; upper apn scales yellowish, lower white; ppn with an exten-

sive area of scales; hypostigial scales present (madrensis) or absent; mep with

flat and semierect scales in a lower anterior angular patch. Legs: forecoxal

Zavortink: Genus Orthopodomyia 109

bristles yellowish-amber, mid- and sometimes hindcoxal bristles darker; fore-

coxa with broad flat scales on anterior surface and above bristles and with semi-

erect scales among bristles, mid- and hindcoxae with broad flat scales in an up-

per outer patch, along outer anterior surface and on midcoxal lower anterior

surface; tarsal segment 1 usually with white scales at base in a small patch or

narrow incomplete or rarely complete ring; hindtarsal segment 5 all white.

Wing: vein Rg 0.91-1.25 length of vein Ro9,3; vein My,9 0.82-1.18 length of

vein M distad of crossvein m-cu; scales symmetrical, moderately broad,

spreading and clasping veins; wing dark, ornamented with light patches (white

to cream or rarely yellowish on basal portion of costa) on costa and other veins,

the number and size of light patches, especially at base of costa, quite variable

within each species, the maximum development as follows: (1) costal vein with

subcostal, apical and usually basal, prehumeral, humeral, presectoral, sec-

toral and accessory subcostal light patches, (2) vein Sc with sectoral, subcos-

tal and sometimes presectoral light patches, (3) vein R with humeral, sectoral,

and sometimes basal light patches, (4) vein R, with accessory sectoral, sub-

costal, apical and usually accessory subcostal light patches, (5) veins R, hans

Cu and sometimes 1A with light patch at base, (6) vein R943 with light natch op-

posite base of vein R445, (7) veins R949 and M with light patch over furcation,

(8) veins M and Cu with light patches distad of crossveins, (9) vein Cu some-

times with light patch basad of its furcation, (10) veins behind R, sometimes

with apical light patch. Abdomen: integument of tergites I-VIII and sternites

II- VIII usually brown, sometimes lighter under areas with light scales; latero-

tergite white-scaled; sternite VIII scaled basally, bare apically.

MALES. Similar to females. Palpus: 0.75-0.98 of proboscis, segment 3

normally without strong apical bristles, usually brown-scaled with light scales

over joints between segments 1-2, 2-3, 3-4 and 4-5 and usually with remainder

of segment 9 light-scaled, light scales white, sometimes yellowish in basal 1

or 2 patches. Antenna: flagellar segment 1 (flavicosta, flavithorax) or segments

1-4 or 5 with tufts of scales. Wing: apical light patches on all or some veins

behind vein Rj sometimes less conspicuous than in female or absent; light patch

over furcation of vein M sometimes reduced.

MALE GENITALIA. Segment VIII: tergite with distinct lobe on posterior

margin. Segment IX: tergite with 0-5 fine bristles on each side. Sidepiece:

long conical; basal mesal lobe usually with 6-8 terete apical bristles, tergal

3-6 quite stout and usually slightly curved mesally at apex, sternal 1-4 usually

more strongly curved mesally at apex and distinctly finer (except in madrensis);

mesal surface completely sclerotized, weakly sclerotized or membranous dis-

tad of basal mesal lobe; mesal surface with patch of specialized curved or sin-

uous bristles distad of basal mesal lobe (albipes, madrensis) or tergomesal

surface with 1 or 2 specialized curved bristles near basal mesal lobe. Spini-

form: normally 1, simple. Aedeagus: weakly to strongly sclerotized, varied

in form and dentition. Proctiger: paraproct sclerotization with 2-5 apical teeth;

cercal setae 1-6.

PUPAE. Cephalothorax: picenia ta varied; integumentary sculpturing

present or absent; hairs largely darker than integument; hairs variously devel-

oped; hair 6-C cephalad of hair 7 and at least hair 7-C strongly developed and

long or hair 6-C caudad of hair 7 (flavicosta, flavithorax); 1 or moreof hairs 1-

9,8,9-C usually strongly developed. Trumpet: largely moderately to strongly

pigmented and darker than darkest part of cephalothorax; shape varied; pinna

varied. Metanotum: pigmentation varied; integumentary sculpturing present

or absent; rarely with fourth pair of hairs developed (flavithorax). Abdomen:

110 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

pigmentation varied; segments I or II-VII or VIII with reticulate integumentary

sculpturing which is moderately to strongly developed on anterior segments but

becomes weaker on posterior segments; smaller hairs concolorous with integ-

ument, larger hairs darker; hair 1-II somewhat dendritic (albipes, wilsoni) or

not, usually 8-28b (4-37); 2-II cephalad of 3-II; 1-VI, VII laterad of hair 2 of

corresponding segment; 9-VII moderately to strongly developed, 7-13b and with

longest branch extending about 0.6-1.1 distance to alveolus of 9-VIII; 9-VIII

strongly developed, usually 11-16b (9-19), and with longest branch reaching

0.4-0.9 distance to apex of paddle. Terminal Segments: male genital lobe var- |

ied. Paddle: lightly pigmented, midrib darker; longer than broad to nearly as

broad as long, shape varied; external buttress nearly straight to convex later-

ally and smooth or with small marginal spicules; midrib nearly straight or con-

vex mesally; entire surface of paddle smooth.

LARVAE. Head: slightly to distinctly broader than long; integument with

or without ornamentation; labrum short; hair 2-C absent. Antenna: short to

moderately long, integument without ornamentation. Thorax: hair 4-M double

or multiple. Abdomen: segment VI without or with (siamensis) a sclerotized

plate, segment VII with or without (flavicosta, flavithorax) a sclerotized plate;

hairs 2-I, II laterad of hair 1 of corresponding segment; 6-I, II multiple, 6-III,

V, VI single, 6-IV often double; 13-I shorter than 10-I; 2-VI, VII mesad of hair

1 of corresponding segment. Segment VIII: sclerotized plate present; comb

scales in 2 rows; posterior row usually with 7-12 (5-14) comb scales; posteri-

or comb scales variously shaped. Siphon: integumentary sculpturing absent or

present. Anal Segment: saddle complete; sclerotized band or incomplete ring

usually present basad of saddle; integumentary sculpturing present or absent;

ventral brush usually with 7 pairs of hairs, the stronger hairs usually with 10-

18 branches.

BIONOMICS. The immature stages are usually found in treeholes, bamboo

internodes or artificial containers. Females of albipes and andamanensis are

known to bite man.

SYSTEMATICS. This is the largest and most diverse group in Orthopodo-

myia, containing 4 subgroups and 9 species, albipes, andamanensis, anopheloi-

des, flavicosta, flavithorax, madrensis, papuensis, siamensis and wilsoni.

The following changes have been made in the (anonomy of this group: maculata

and maculipes, considered to be distinct taxa by Knight and Mattingly (1950:9,

10), and lemmonae, described by Thurman (1959:58-59), are here considered

as synonyms of anopheloides; mcgregori, a name applied to a Philippine speci-

men by Knight and Chamberlain (1948: 10) and Knight and Mattingly (1950:13-15),

is treated as a nomen dubium; and papuensis and siamensis are described as

new species. | |

Knight and Mattingly (1950:2) placed flavicosta and flavithorax with the

Mauritian arboricollis and the Madagascan vernoni on adult features. After a

consideration of the immature stages I cannot agree and believe that flavicosta ©

and flavithorax should be allied with the species which these authors placed in

the anopheloides subgroup. The Albipes group of this paper is then equivalent

to the anopheloides subgroup of Knight and Mattingly plus flavicosta and flavi-

thorax. As indicated below, the Anopheloides subgroup is here restricted to

the 3 species anopheloides, andamanensis and papuensis.

The greatest discordance among classifications based on immature stages,

male genitalia and adults is found in this group. Based on adult characters,

flavicosta and flavithorax would form one group while the other 7 species would

form a Second; of the latter 7 species, albipes, andamanensis, anopheloides,

Zavortink: Genus Orthopodomyia kot

papuensis and siamensis are very Similar while madrensis and wilsoni each dif-

fer slightly. Similarities in male genitalia would group andamanensis and anoph-

eloides, place albipes, flavicosta, flavithorax and wilsoni together, aggregate

papuensis and siamensis,’ and put madrensis into a separate category. Groups

based on pupae would be (1) flavicosta, flavithorax, madrensis and siamensis,

(2) wilsoni, (3) albipes, and (4) papuensis, andamanensis and anopheloides. The

larval classification agrees with the pupal except that flavithorax would form a

fifth group. The classification used here is the one shown by the pupae and the

group is accordingly divided into the Flavicosta, Wilsoni, Albipes and Anophe-

loides subgroups.

Hybridization may explain at least part of the discordance between the

stages. The Anopheloides subgroup may be of hybrid origin, with putative an-

cestors in the Albipes and Flavicosta subgroups.

In light of the great variety of larval and pupal types, the Albipes group is

probably one of the oldest in the genus. Its relationships have not been deter-

mined. The larval chaetotaxy of wilsoni is very similar to the Central Ameri-

can phyllozoa whereas the larval chaetotaxy of the Anopheloides subgroup is

similar to the Holarctic Signifera group. Adult mesonotal ornamentation is like

the Mauritian arboricollis, the Madagascan Vernoni group, and the South Amer-

ican Thomasina section, while the ornamentation of the wing is like phyllozoa.

The male genitalia of several species have a more or less pyriform aedeagus

which is Somewhat similar to the Signifera group and arboricollis.

Adults of most species in this group are rather variable, particularly in

ornamentation of the proboscis, wing, abdomen and hindtarsus. The larval

chaetotaxy of albipes, anopheloides and wilsoni is rather variable and hairy

forms are known for the larva and pupa of anopheloides and the pupae of anda-

manensis and papuensis. These hairy forms resemble the normal larva and

pupa of albipes in many ways.

Identification of male genitalia is difficult because many species differ only

subtly in the shape of the aedeagus. It is necessary to have the genitalia re-

moved from the eighth segment in order to see the often weakly sclerotized ae-

deagus, and in addition the genitalia must be mounted perfectly horizontally and

with the parts in the normal resting position.

The group differs from the others in the genus in that every species except

papuensis is sympatric with at least 1 other species over all or part of its range.

Anopheloides occurs with every species in the group except papuensis and in

some areas 3, 4 (Malaya) or possibly even 5 (Thailand) species may be sym-

patric: :

DISTRIBUTION. India, southern China and southern Japan, south to Cey-

lon, the Malay Archipelago and northern Queensland.

KEYS TO SPECIES

Adults

1. Femora, tibiae and tarsal segments 1 with light scales aggregated into

distinct light patches; costal vein and vein Rj lacking a white patch

opposite furcation of vein R943 into veins Ro and Rg (accessory sub-

costal patch); paratergite with a small patch of light scales. . . . 2

Femora, tibiae and tarsal segments 1 with light scales largely scattered,

costal vein and vein Ry normally with an accessory subcostal light

patch; paratercite bare ia te ei tw ee i 8

112

2(1).

3(1).

4(3).

5(4).

6(5).

7(6).

8(7).

att}.

Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

Mesonotal-scaling largely bright yellow or golden; costal vein with pre-

sectoral light patch (at least ventrally) and at most with only a few

light speckles in basal 0.5; light-scaling on ventral surface of pro-

boscis of female distad of light patch on dorsal surface; vein Cu with

or without a light patch basad of its furcation in female 17. flavithorax

Mesonotal-scaling appearing largely brownish; costal vein without pre-

sectoral light patch and with numerous light speckles in basal 0.5;

light-scaling on ventral surface of proboscis of female basad of light

patch on dorsal surface; vein Cu always with a light patch basad of its

furcation in feniale. 6 owiite es ewe aly Leto telus wee fievicosta

Pra with 1 bristle; hypostigial scales present ... . . 18. madrensis

Pra normally without a bristle; hypostigial scales absent ...... 4

Hindtarsal segments 2 and 3 with a moderate-sized subequal white ring

at each end; tergites II and sometimes III of female with a large mid-

dorsal basal light (usually yellowish) patch . ... . . . 20. wilsoni

Hindtarsal segments 2 and 3 not as above, segment 2 entirely or largely

dark-scaled at either base or apex, segment 3 all white or with dark

scales in proximal or distal portion; tergites II and III of female with-

outa large middorsal:basal light patel. vies 6.6 a owees chcwre “5

Hindtarsal segment 2 usually with more extensive white-scaling at base

than at apex; palpus of female with three white patches beyond the ©

base; abdominal sternites II or il to V or VI with a midventral apical

light paten oGe fei . oe! won ede albipes

Hindtarsal segment 2 always mallet more sentative wiles ~gealine at apex

than at base; palpus of female with two white patches beyond the base;

abdominal sternites II-VI without a midventral apical light patch. . 6

Hindtarsal segment 4 with a dark spot or ring distad of middle. .. .

24, in ps anopheloides

Hindtarsal comment 4 oie uglite~wealbdli

Hindtarsal segment 3 entirely white-scaled. . . .19. in part, siamensis;

22. papuensis; 23. in part, andamanensis; 24. in part, anopheloides

Hindtarsal segment 3 with some dark scales . . 0.0. 6 ee wee ee B

Hindtarsal segment 3 with dark-scaling in basal 0.5. eae

. OBS. im part, andavhariensis

Hindtarsal seement 3 with dark- scaling in n apical 0.5 pve Miah

. 19. in part, siamensis; 24. in part, aicohmaider

Male Genitalia

Aedeagus without any sclerotized teeth on the sternal surface but with a

projection from the base which extends between or below the ventral

parameres. . . 2

Aedeagus with one sclévovimed tooth ¢ on ‘phen half el sietiul eurtace and

without a projection extending between or below the ventral para-

GEOR aan gee a Gy eae ae EE eS eS 8

Basal projection of aedeagus usually extending conspicuously between the

Sit).

4(3).

5(4).

6(4).

7(6).

8(6).

Dt).

Zavortink: Genus Orthopodomyia 113

ventral parameres . . . . . .'. 24, anopheloides

Basal projection of aedeagus usually ‘dispated more sternally so that it

does not extend CORBPIORONE between the ventral parameres .

ee andamanensis

Basal mesal lobe without apical bristles sharply divided into strong and

fine ones, but with 7 or 8 more or less equally developed, strong

bristles; mesal surface of sidepiece with a large patch of strong, sin-

uous, specialized bristles distad of basal mesal lobe . .18. madrensis

Basal mesal lobe with 3-6 strong apical bristles and 1-4 finer ones;

mesal surface of sidepiece with or without a patch of strong sinuous

or curved specialized bristles distad of basal mesal lobe . .... 4

Aedeagus not distinctly pyriform, broadest in apical halt or at or distad

of level of sternal teeth... . 2

Aedeagus more or less pyriform, distinctly broadest i in basal half and

basad of level of stermal (6G eo ee a aa

Sternal aedeagal teeth 0. 50-0.55 distance from base and usually basad

of broadest portion of aedeagus. . . . .19. siamensis

Sternal aedeagal teeth 0.65-0.75 distance Eom base and usually slightly

distad of broadest portion of aedeagus. . .... . . . 22. papuensis -

Sclerotized teeth on sternal surface of aedeagus separated by a distance

nearly equal to the width of one of them; tergite VIII lobe with numer-

ous distinct serrations at apex. . . 7

Sclerotized teeth on sternal surface of aedeagus aandienoue or r nearly So;

tergite VIII lobe without serrations at apex or with only a few irregu-

Tae LOC iat oe ee rer eed ee eh a a ee ee ee B

Mesal surface of sidepiece with a large patch of strong, sinuous or

curved, specialized bristles distad of basal mesal lobe. . 21. albipes

Mesal surface of sidepiece without specialized bristles . . . 20. wilsoni

Clasper strongly curved in the distal 0.25; each strong bristle of basal

mesal lobe conspicuously mane than the one dorsad of it. .

«eh OE flavicosta

Clasper ae slightly curved in distal 0. 25: “each strone bristle of basal

mesal lobe only slightly longer than the onedorsad ofit 17. flavithorax

Pupae

Trumpet cylindrical and largely dark brown; hairs 1-6, 8,9-C moderate-

ly developed, shorter than the trumpet, and usually 1-3b (1-4)... .

‘ .(WILSONI SUBGROUP) 20. wilsoni

Trampet not both cylindrical and dark brown; normally one, some, or

most of hairs 1-6, 8,9-C strongly to very reset developed, longer |

than the trumpet, wae or with more than 4b. . 2

Trumpet attaining maximum or nearly maximum width in basal 0.33, the

remainder broad and almost or actually parallel sided . .

5 (ALBIPES SUBGROUP) 21. ‘albipes

Trumpet bacadenine sradually from base to apex, or if attaining nearly

114

S{2).

4(3).

(4).

6(5).

7(3).

8(7).

att):

Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

maximum width in basal 0.5, trumpet is rather narrow. ..... 3

Hair 5-VI long, extending to or beyond alveolus of hair 4-VIII, and sin-

gle, or if 2,3b, then 6-C is caudad of 7-C and 5-II is short and multi-

ple. : . (FLAVICOSTA SUBGROUP) 4

Hair 5-VI short, "extending nearly to or - slightly beyond alveolus of 4-VII,

or mies? usually ob; 6=C pesataren of 7-C and 5-II long and single .

‘ : (ANOPHELOIDES SUBGROUP) 7

Hair 5=Vi 2, 5b; 5-Tl short, 3b; metanotum with 4 pairs of hairs .

yond 17. flavithorax

fair 3 -VI single; at Tl tone single: ieradotiin wie 3 pairs of hairs . 5

Hairs 1, 2-C strongly developed and usually 6, 7b (5-8); 2-II laterad of

5-II, 2-IV thickened. . . . .19. siamensis

Hairs 1 2-C moderately developed and usually 1 2 3b a- ” 2-II mesad of

ey Dee Aae CO 8 a

Hair 6-C caudad of 7-C; 5-III 4-6b; 5-IV single ... . . 16. flavicosta

Hair 6-C hig eet of T- c 5-III 2, 3b; 5-IV See | 3b (2- 4). :

ees, ‘ : 18. Wieeroneis

Hairs 5-V, VI long, extending to or nearly to alveolus of fair 4 of second

following segment; usually some or all of hairs 2-4-C and 6-III, V, VI

very strongly developed and long .

. (hairy forms) 22. papuensis; 23. aida anensis: 24, anopheloides

Hairs a -V,VI short, extending nearly to or slightly beyond alveolus of

hair 4 of first following segment; usually none of hairs 2-4-C or 6-

III, V, VI more than moderately developed . . . . (non-hairy forms) 8

Hair 1-III usually 12-14b (10- all 9-C aswally 2, 3b (2-5) :

. . 22. papuensis; 23. waaianensic

ae 1-0 sively 5- Tb (3-11); ge C oe a (single, double).

Ce a ee a : : : D4, anophelowes

Larvae

Hairs 4-10-C all more or less equally developed, long, strong and mul-

tiple; 0,1,3-7-P all more or less equally developed, long, strong and

multiple. . . . . (FLAVICOSTA SUBGROUP, in part) 17. flavithorax

Hairs 4-10-C not all equally developed, in particular 8-10-C only 1-3b;

0,1,3-7-P not all Meu rar cae in particular 1,5,6-P normally

nate te SPO pre ee Dh re eg te

Hair 1-M shorter than or only slightly longer than 3-M; 1-T shorter

than or only slightly longer than 2-T; posterior comb scales with 9-7

large apical spines; 6-I, II with individual branches usually stouter

tran Ode ae ; (FLAVICOSTA SUBGROUP, in part) 3

Not with above Sonibianes of characters; if 1,3-M and/or 1, 2- T are

Subequal in length, then posterior Corb scales have only a single ter-

minal spine, or if posterior comb scales have more than 1 terminal

spine, then 1-M is much longer than 3-M and 1-T is much longer than

2-T; 6-I, II with individual branches usually finer than 6-TT.... 9

Zavortink: Genus Orthopodomyia — EE

3(2). Hairs 1-M, T multiple; 1-III-V shorter than or subequal to hair 3 of cor-

responding segment; head capsule papillose. . .. . .16. flavicosta

Hairs 1-M, T single; 1-III-V eer than hair 3 of 6 Aduwon & seg-

ment; head capsule smooth ey ; ;

4(3). Hairs 6-I, II with branches coarse nearly to apex and barbed only in bas-

al portion; 13-C usually 3-5b; 1-V usually single .. 18. madrensis

Hairs 6-I, II with branches tapering to a fine apex and barbed from near

base ka near apex; 13-C ape be 8, 9b a ae 1-V cree double .

: . : .19. siamensis

5(2). Longest spine of posterior comb scales not longer than 1.5 times the

distance from its base to the attachment of the comb scale . .

; (ALBIPES SUBGROUP) 21. albipes

Longest some or postoniar eprats scales longer than 2.0 times the dis-

tance from its base to the attachment of the comb scale. . .... 6

6(5). Posterior comb scales with only 1 large apical spine . .

; (WILSONI SUBGROUP) 20. ‘wilsoni

Posterior ‘conb scales oa 3- 5 panels apical spines. . .

. (ANOPHELOIDES SUBGROUP) 7

7(6). Most or all of hairs 1-M, T, III, V multiple; 1-VI often longer than 4-VI .

(hairy forms) 24. anopheloides

All ol hairs Ie -M, qT, Il, Vv usually singles bay 1-VI ee shorter than

BV a. ee ee . (non-hairy forms) 8

8(7). Hair 6-1 usually 9, 10b (7-10); hair 6-II pi ie 9-11b (7-12) .

eos ani boneume

Hair 6- I {usually 5- 8b; hair 6- I usually 4- 8b .

wee: papuensis: 24. anopelcides

FLAVICOSTA SUBGROUP

PUPAE. Abdomen: hair 1-II strongly developed and long and with mostly

simple branches, some much finer than others, arising from the base; 5-V, VI

long, extending nearly to or beyond alveolus of hair 4 of second following seg-

ment; 5-VI usually single or double (1-3b). Terminal Segments: male genital

lobe projecting far beyond median caudal lobe. Paddle: asymmetrically obo-

vate in outline.

LARVAE. (Description does not pertain to the highly modified flavithorax).

Head: distinctly broader than long; hairs 4-10-C not all equally developed.

Thorax: hairs 0,1,3-7-P not all equally developed; 1-M, T short; 13-T weakly

developed, usually 3-5b. Abdomen: hairs 6-I, II with individual branches stout-

er than 6-III. Segment VIII: posterior comb scales with 5-7 large apical

spines. |

BIONOMICS. Larvae have usually been found in treeholes. Habits of the

adults are unknown.

SYSTEMATICS. This subgroup contains the 4 species flavicosta, flavitho-

rax, madrensis and siamensis. The 3 allopatric species flavicosta, madrensis

and Siamensis are well marked from each other as adults, male genitalia, pu-

116 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

pae and larvae, yet clearly belong to 1 group on larval and pupal characters.

The fourth species, flavithorax, is sympatric with flavicosta; its larva is very

different from that of the other 3 species and shows no evidence of belonging to

the same subgroup, but its pupa, while showing several unique features, is

Similar to the other species and the adult and male genitalia resemble flavicos-

ta. There appear to be no characters in the adults or male genitalia to unite

these 4 species into 1 subgroup, although 3 of them, flavicosta, flavithorax and

madrensis, are the only members of the Albipes group to have a bristle on pra.

Since 3 of the 4 species are well marked in all stages, and all 4 species

have very restricted, relictual distributions, this must be one of the oldest

subgroups in the Albipes group. :

DISTRIBUTION. Discontinuous, with species found in southwestern India

and Ceylon, peninsular Thailand and the Philippines.

16. Orthopodomyia flavicosta Barraud

Figs. 23, 24

1927. Orthopodomyia flavicosta Barraud, 1927:531-532. *TYPE: lectotype ¢

(875) with genitalia slide 660610-4, specimen labeled ''TYPE o,'' Karwar,

[Mysore], India, reared from a larva found in a treehole, Sept 1921,

P.J. Barraud; PRESENT SELECTION [BMNH].

Orthopodomyia flavicosta of Barraud and Covell (1928:676); Barraud (1932:1016);

Edwards (1932:108); Barraud (1934:104-105); Knight and Mattingly (1950:2);

Stone, Knight and Starcke (1959:123).

FEMALE. Wing: 3.40 mm. Proboscis: 2.02 mm. Forefemur: 2.24 mm.

Abdomen: about 2.5 mm. Head: integument of head capsule, clypeus, torus

and ends of most flagellar segments dark brown, middle portion of most flagel-

lar segments lighter; frontal and orbital bristles amber to brown; decumbent

scales silvery-white to cream-colored; erect scales short, broad and dingy

yellowish-brown to brown posteriorly and somewhat longer, narrower and light-

er anteriorly, a few erect scales, especially laterally, white or yellow; dorso-

lateral scales brown anteriorly, white posteriorly; lateral and ventral scales

white; labium mostly brown-scaled, white scales scattered ventrally near base

and in dorsal patch about 0.67-0.75 distance from base, white and/or yellow

scales in ventral patch basad of dorsal white patch, and near apex dorsally;

palpus 0.51 of proboscis, 4-segmented, segment 4 short, mostly brown-scaled,

white scales over joint between segments 2 and 3 and covering all of segment 4

and sometimes in a patch near base of segment 2; torus scales white; flagellar

scales white and brown. Thorax: integument of mesonotum, postnotum and

pleuron uniformly brown to dark brown, scutellum lighter; paratergite with a

few dingy to white scales; acrostichal scales brown at caudal end, remainder

brown and golden or mostly golden; dorsocentral scales white or golden at cau-

dal end, remainder brown and golden; lateral prescutellar scales golden or

brown and golden, with white scales laterally and caudally; posterior fossal

scales mostly golden or yellow; fossal scales usually largely brown anteriorly

and golden or yellow posteriorly; supraalar scales white mesally and in anteri-

or projection, remainder golden or yellow; pra with I bristle; mep bristles

yellowish amber; ppn with white and yellow decumbent scales along upper mar-

gin, yellow decumbent scales along posterior margin and flat white scales along

Zavortink: Genus Orthopodomyia 117

ventral margin; pra, ppl and pst scales white or yellow-tinged; ssp scales ina

long row, white anterioriy, yellow-tinged posteriorly; hypostigial scales absent;

upper half of stp with bare areas near lower end of ssp scales and opposite up-

per end and middle of lower mep scale patch, upper and anterior scales white,

remainder yellow-tinged; lower stp scales yellow-tinged above, white below;

lower mep scales white. Legs: integument of mid- and hindcoxae same color

as pleuron, forecoxa sometimes lighter; coxal scales white except for a few

yellowish ones ventrally on forecoxa and some brown ones along outer anterior

edge of mid- and hindcoxae; femora brown-scaled with irregular white to beige

patches on anterior surface and on posterior surface of forefemur, posterior

surface of mid- and hindfemora with pale speckles and usually with a pale-

scaled streak or patch at least near apex; tibiae brown-scaled with many irreg- .

ular white patches on all surfaces (each patch is distad of a bristle); tarsi

largely brown-scaled, segment 1 with small patches of white scales; foretarsal

segment 2 and sometimes segment 3 with dorsobasal light patch; midtarsal seg-

ment 1 with a few light scales at apex, segments 2,3 and sometimes 4 with dor-

sobasal light patch; hindtarsus with white rings over joints between segments

1-2, 2-3 and 3-4 and with white scales at apex of segment 4. Wing: costal vein

without presectoral, sectoral or accessory subcostal patches, with or without a

single irregular patch in prehumeral to humeral area, and with numerous yel-

low to white scales speckled from the basal to the subcostal patch and usually

at position of accessory subcostal spot; vein Sc with sectoral patch sometimes

very small, presectoral patch absent; vein R with basal patch; vein Rj without

accessory subcostal patch but usually with light speckles in that area; vein 1A

dark at base; vein Cu with light patch basad of its furcation; usually all veins

behind Ry, except sometimes Cug, with apical light patch. Haltere: integument

beige to brown; scales transparent to brownish. Abdomen: tergite I with brown-

scaled fan and scattered brown and white scales laterally; tergites II-VII large-

ly brown-scaled, with basolateral white patch and smaller apicolateral white to

yellowish patch which becomes even smaller or disappears on distal segments;

tergites III-VII with subdorsal median yellowish or white and yellowish patch;

tergite VIII brown-scaled with basal white band which is broader laterally; ster-

nites II-VII with basolateral white patches which are large and cover most of

sternites II and III but become smaller on distal segments, remainder brown-

scaled; all or only more distal of sternites II-VI sometimes with ventral medi-

an light patch; sternite VIII scales dark.

MALE. Similar to female. Proboscis: white scales in narrow incomplete

to complete ring just distad of level of distal end of palpal segment 2 and usual-

ly speckled or streaked for a short distance distad of this ring on ventral sur-

face, yellow scales in dorsal preapical patch and scattered, especially in api-

cal portion. Palpus: about 0.94 of proboscis; segment 3 with yellow speckles.

Antenna: flagellar segment 1 with tuft of large and small brown and white scales.

Abdomen: sternite VIII dark-scaled with light scales apically; sidepiece with

light and dark scales. |

MALE GENITALIA (fig. 23). Segment VIII: tergite lobe much longer than

broad, wider apically than basally, apex truncate, without teeth. Segment IX:

tergite with 2-5 bristles on each side. Sidepiece: basal mesal lobe with 3 or

4 stout bristles and 2-4 finer; each stout bristle considerably longer than the

one dorsad of it; mesal surface membranous distad of basal mesal lobe; spe-

cialized mesal bristles absent, 1 or 2 specialized tergomesal bristles present.

Clasper: strongly curved in distal 0.25. Aedeagus: moderately sclerotized;

more or less pyriform, with 1 pair of contiguous or nearly contiguous sternal

118 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

teeth about 0.65 from base; without basal projection between parameres. Proc-

tiger: paraproct sclerotization usually with 3 or 4 apical teeth; cercal setae 2

or 3.

PUPA (fig. 23). Abdomen: 3.22 mm. Trumpet: 0.40 mm. Paddle: 0.73

mm. Cephalothorax: mostly light greyish-tan colored, posterior portion of

mesonotum and upper part of wing case darker, tan; integumentary sculpturing

conspicuous, reticulate; hairs 1,2-C moderately developed, usually double (2,

3b); 3-C moderately developed, 3-6b; 4-C moderately developed, 2-4b; 5-C

with 1 branch very strongly developed, others moderately developed, 2-4b; 6-

C caudad of 7-C; 7-C very strongly developed, double; 8-C moderately devel-

oped, single; 9-C moderately developed, double. Trumpet: light brown in color

except for pinna which is lighter; widening gradually from base; pinna large.

Metanotum: light brown mesally, light greyish-tan laterally and on haltere case;

reticulate integumentary sculpturing well developed; hair 10-C moderately de-

veloped, 5-9b; 11-C moderately developed, usually with 2, 3b arising below mid-

dle; 12-C moderately developed, 3,4b; fourth pair of hairs never developed.

Abdomen: anterior segments light tan, posterior very light greyish-tan; poste-

rior edges of anterior segments darker; integumentary sculpturing strongly de-

veloped on anterior segments; hair 1-II usually 21-27b; 2-II slightly thickened

and mesad of 5-II; 5-II long and single; 2-III slightly thickened, 2-IV fine; 5-

IV-VI single; 9-VII 9-12b and with longest branch extending to alveolus of 9-VIII;

9-VIII 9-14b and with longest branch reaching about 0.6-0.7 distance to apex of

paddle. Paddle: very light straw-colored; quite angular in outline, with great-

- est width nearly equal to length; external buttress slightly convex laterally in

distal portion and with small marginal spicules; midrib nearly straight to slight-

ly convex meSally. :

LARVA (fig. 24). Head: 1.08 mm. Siphon: 1.23 mm. Anal Segment: 0.31

mm. Head: integumentary sculpturing conspicuous, papillose; largely straw-col-

ored, becoming darker, tan or brown, posteriorly, collar dark brown; mental

plate straw to tan, usually with 9,10 teeth on each side; 13-C weakly developed,

usually 2,3b. Antenna: moderately long, tan to light brown basally, becoming

straw-colored apically. Thorax: epidermal pigment absent; spicules absent;

8-P usually 2, 3b; 1-M usually 2-4b; 1-T usually 3-6b. Abdomen: segment VII

without a plate; hairs 1-I-IV short, usually 3, 4b (3-5); 6-I, II with individual

branches tapering to a fine apex and barbed from near base to near apex; 1-V

and 13-III-V short, 2,3b. Segment VIII: sclerotized plate large but not ringing

segment; anterior comb scales usually 21-24 (20-25), posterior comb scales

usually 7-10 (7-11); anterior comb scales fringed only; posterior comb scales

usually with 5 large spines and 2 smaller ones; posterior row of comb scales

about 0.65-0. 84 length of anterior row; hair 3-VIII usually 6, 7b. Siphon: index

about 4.5-5.7; base of siphon brown to dark brown, portion between base and

alveolus of hair 1-S tan to brown, portion between 1-S and apex darker, dark

tan to dark brown, apex straw-colored; strongly developed imbricate integu-

mentary sculpturing present; hair 1-S located 0.40-0.44 from base of siphon,

usually 5-8b (4-10). Anal Segment: base and dorsal surface brown, remainder

tan to light brown; moderately developed imbricate integumentary sculpturing

present; dorsal anal gills about 1.5-1.6 length of ventral pair and about 0. 7-

0.8 length of anal saddle.

BIONOMICS. The immature stages have been found only in treeholes and

may occur in the same breeding site with flavithorax and anopheloides.

SYSTEMATICS. Adults of this species and the following, flavithorax, are

most conspicuously differentiated from all other members of the Albipes group

Zavortink: Genus Orthopodomyia : | 119

by the following features: (1) definite patches rather than scattered light scales

on the femora, tibiae and first tarsal segments, (2) absence of the accessory

subcostal light patch on the costal vein and vein Ry; (3) a light patch basad of

the fork of vein Cu (at least in the male), (4) a broad white ring over the joint

between hindtarsal segments 1 and 2, (5) a small patch of scales on the para-

tergite, (6) scales on only the first flagellar segment in the male, and (7) the

male palpus about as long as the proboscis. The adults of flavicosta and flavi-

thorax are distinguished from each other by the characters given in the key.

The ornamentation of the basal portion of the costal vein is quite variable

in flavicosta. There may or may not be a small to large light patch in the pre-

humeral to humeral area, but there is never a presectoral patch. There are

numerous yellow, or rarely white, scales scattered along the basal half of the

costa. According to Barraud (1927:532) these scales may be so numerous as

to form a long streak.

The male genitalia of flavicosta and flavithorax are quite distinct from those

of the other 2 species in the Flavicosta subgroup, but resemble those of alb ipes

and wilsoni. The pupal chaetotaxy of flavicosta is more similar to madrensis

than any other species in this subgroup, but it shares with flavithorax the place-

ment of hair 6-C caudad of 7-C. The larva of flavicosta is not as similar to

either madrensis or siamensis as those two species are to each other.

One of Barraud's specimens from Karwar, India, cannot be assigned to

either flavicosta or flavithorax and is apparently a hybrid between the 2 species.

The larval skin is in such poor condition that few hairs can be checked, but the

skin is definitely not the extremely hairy flavithorax and agrees with flavicosta

in comb scales and those hairs which can be identified. The pupa, adult, and

male genitalia show mixed flavicosta and flavithorax characters.

Flavicosta

DISTRIBUTION. Coastal southwestern India. Material Examined: 36 spec-

imens; 7°, 92, 3P, 17 L; 3 larval individual rearings.

; INDIA. Mysore: Karwar, Aug-Sept 1921, P.J. Barraud, & lectotype (875),

2160 (876,886): 1 1p24873), 26. 6o 14:L ‘BMNH], 2%, 22 [USNM].

Additional records from the literature. None.

Hybrids

DISTRIBUTION. Karwar, India. Material Examined: 3 specimens; 1 <,

1 P, 1 L; 1 larval individual rearing.

INDIA. Mysore: Karwar, Aug 1921, P.J. Barraud, 1 lpo (882).

Additional records from the literature. None.

17. Orthopodomyia flavithorax Barraud

Figs. 25, 26

1927. Orthopodomyia flavithorax Barraud, 1927:529-531. *TYPE: lectotype “

(871) with genitalia slide 660610-2, specimen labeled ''Type o,'' Karwar,

[Mysore], India, reared from a larva found in a treehole, Sept 1921,

P.J. Barraud; PRESENT SELECTION [BMNH].

Orthopodomyia flavithorax of Barraud and Covell (1928:676); Barraud (1932:

1015-1016); Edwards (1932:107, 108); Barraud (1934:103-104); Knight and

120 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

Mattingly (1950:2); Stone, Knight and Starcke (1959:123).

FEMALE. Wing: 3.63 mm. Proboscis: 1.98 mm. Forefemur: 2.05 mm.

Abdomen: about 2.5mm. Very similar to flavicosta, differing in the following

details. Head: decumbent scales yellow except for white ones at base of fron-

tal bristles and in front of dorsolateral scales; many erect scales, especially

laterally, yellow; labium with dorsal white patch about 0.67-0.75 distance from

base and with white and/or yellow scales in ventral patch distad of dorsal white

patch and sometimes also scattered near base ventrally and apex dorsally or

ventrally. Thorax: acrostichal scales brown at caudal end, remainder bright

yellow; dorsocentral scales bright yellow; lateral prescutellar scales brown

and bright yellow; posterior fossal scales bright yellow; fossal scales yellow-

ish-brown in center of fossa, remainder bright yellow; supraalar scales white

in anterior projection, remainder bright yellow; all ppn scales curved, decum-

bent, yellow along upper and posterior margins, white along ventral margin;

pra scales yellowish; ssp scales mostly yellow-tinged; stp scales white anteri-

orly, remainder yellow-tinged. Legs: forecoxal scales all white; posterior

surface of mid- and hindfemora with more extensive pale-scaling, sometimes

largely yellowish; midtarsal segment 1 without white scales at apex. Wing:

costal vein with presectoral patch present at least on ventral surface (usually

present on dorsal surface also), always with a single patch in prehumeral to

humeral area, and with or without a few yellow scales speckled from the pre-

sectoral to the subcostal patch; vein Cu with or without light patch basad of its

furcation. Abdomen: all or only more distal of tergites II-VII with subdorsal

median white and/or yellowish patch; basolateral light patches on distal ster-

nites yellowish and sometimes connected to form basal bands; sternite ok all

dark-scaled basally or with basolateral light patch.

MALE. Similar to female; differs from flavicosta as follows. Proboscis:

with or without small patch or narrow ring distad of level of distal end of pal-

pal segment 2, without distinct yellow preapical patch, with many yellow and

white scales speckled or arranged into small spots throughout length but espe-

cially numerous apically. Palpus: about 0.98 of proboscis. Wing: vein Cu al-

ways with light patch basad of its furcation (always so in flavicosta o and @ but

not always so in flavithorax 2). Abdomen: sternite VIII light-scaled basally,

laterally and apically.

MALE GENITALIA (fig. 25). Segment VIII: tergite lobe not or considerably

longer than broad, sides parallel or diverging apically, apex truncate or emar-

ginate, without distinct serrations. Segment IX: tergite with 0-3 bristles on

each side. Sidepiece: basal mesal lobe with 4 or 5 stout bristles and 1 or 2

finer; each stout bristle only slightly longer than the 1 dorsad of it; mesal sur-

face membranous distad of basal mesal lobe; specialized mesal bristles absent,

1 or 2 specialized tergomesal bristles present. Clasper: not strongly curved

in distal 0.25. Aedeagus: moderately sclerotized; more or less pyriform, with

1 pair of contiguous or nearly contiguous sternal teeth about 0.5-0.7 from base;

without basal projection between parameres. Proctiger: paraproct sclerotiza-

tion usually with 3 or 4 apical teeth; cercal setae 1-3.

PUPA (fig.25). Abdomen: 3.36 mm. Trumpet: 0.46 mm. Paddle: 0.70

mm. Cephalothorax: ventral portion light greyish-tan, dorsal portion tan, pos-

terior part of mesonotum and upper part of wing case light brown; integumen-

tary sculpturing absent; hair 1-C strongly developed, usually double (2, 3b);

hair 2-C strongly developed, 6-12b; hair 3-C strongly developed, 5-9b; hair

4-C moderately developed, 4, 5b; 5-C strongly developed, 5-7b; 6-C caudad of

Zavortink: Genus Orthopodomyia 121

T-C; 7-C strongly developed, 4, 5b; 8-C strongly developed, 3b; 9-C strongly

developed, 7,8b. Trumpet: mostly brown in color, extreme base and apex

lighter; broadened gradually from base; pinna large. Metanotum: evenly light

brown; integumentary sculpturing absent; hair 10-C moderately developed, 14-

21b; 11-C moderately developed and with 4-6b from near base; 12-C strongly

developed, 9-11b; fourth pair of hairs developed, moderate, 3,4b. Abdomen:

anterior segments evenly light brown, posterior segments straw- colored; in-

tegumentary sculpturing strongly developed on anterior segments; hair 1-II

16-21b; 2-II slightly thickened and mesad of 5-II; 5-II short, 3b; 2-III, IV thick-

ened; 5-IV, V 3b; 5-VI usually double (2, 3b); 9-VII 8b and wath longest brafich

extending beyond alveolus of 9-VIII; 9-VIII 11,12b and with longest branch

reaching 0.6-0.7 distance to apex of paddle. Paddle: light straw-colored; sim-

ilar to flavicosta, but not as angular; external buttress convex laterally and :

smooth; midrib slightly convex mesally.

LARVA (fig.26). Head: 1.12 mm. Siphon: 1.50 mm. Anal Segment: 0.39

mm. Head: distinctly broader than long; integumentary sculpturing absent;

largely straw-colored to light tan, lighter around imaginal eye, darker poste-

riorly, collar brown; mental plate straw-colored to tan, usually with 7, 8 (7-9)

teeth on each side; hairs 4-10-C all more or less equally developed, strong,

long and usually 12- 18b (8-22); 13-C strongly developed, 8-12b. Antenna: mod-

erately long, base brown, becoming straw-colored apically. Thorax: epider-

mal pigment absent; spicules present; hairs 0,1,3-7-P all more or less equal-

ly developed, strong, long and usually 8-24b (6- 27): 8-P usually (?) 14-18b;

1-M long, usually 26-30b; 1-T long, usually 15-23b; 13-T strongly developed,

usually 5,6b. Abdomen: segment VII without plate; hair 1-I moderately long,

7-9b; 1-II-IV moderately long, usually 4-6b; 6-I, II with individual branches

finer than 6-III, tapering to a fine apex, and barbed from near base to near

apex; 13-III-V moderately long, 6-9b; 1-V long, 3-5b. Segment VIII: sclero-

tized plate present, but reduced and irregular in all specimens available for

study; anterior comb scales usually 19-22 (17-26), posterior comb scales usu-

ally 10-12; anterior comb scales fringed; posterior comb scales with 1 very

long, strong apical spine and 3 or 4 smaller spines on each side of base of large

Spine; posterior row of comb scales 0. 89-1.02 length of anterior row; hair 3-

VIII usually 5, 6b. Siphon: index about 5.3-6.3; colored as in flavicosta; in-

tegumentary sculpturing imbricate, moderately developed basally; hair 1-S lo-

cated 0.39-0.42 from base of siphon and usually 6-9b. Anal Segment: colora-

tion and sculpturing as in flavicosta; dorsal anal gills about 1.3-1.7 times

length of ventral pair and about 0.3-0.6 length of anal saddle.

BIONOMICS. Larvae have been collected only in treeholes, where iney

may be associated with those of flavicosta and anopheloides.

SYSTEMATICS. The adult of this species is very similar to flavicosta and

differs from the remaining species in the Albipes group in the same features.

The scaling at the base of the costal vein is less variable in this species

than in flavicosta. There are always 2 light patches, one in the prehumeral to

humeral area, and one in the presectoral area. The latter patch is represen-

ted on the upper surface of the wing in all specimens except one and in that

specimen it is present on the ventral surface. The ornamentation of vein Cu

is more variable in the female of this species than in flavicosta, since a light

patch may be present or absent basad of the furcation of the vein.

The pupa of flavithorax is more similar to siamensis than to any other spe-

cies, but it has in common with flavicosta the position of hair 6-C caudad of 7-

C. The development of the fourth pair of hairs on the metanotum of this species

122 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

is unique in the genus. Only 2 specimens of the pupa have been available for |

study, but since the hair is present and well developed on both sides of each.

individual I think that it must be a normal component of the chaetotaxy rather

than an anomaly. |

The flavithorax larva. is completely unlike that of any other in the Flavicos-

ta subgroup and is, in fact, the most distinctive larva in the entire genus. It is

remarkable in the elongation and branching of many hairs on the head and tho-

rax. Even the comb scales are of a type not found elsewhere in the Albipes

group. The extremely different larva of this species poses problems in erecting

a classification. The species could actually belong to a separate subgroup and

have obtained adult and male genitalic features similar to those of flavicosta via

introgression with that species, or the species could truly be related to flavi-

costa and the peculiar larva have been evolved in response to competition with

it. The second alternative has been chosen here.

Barraud (1932:1013) stated that no definite saddle occurred on segment VIII

of the larva, but that there was some indefinite sclerotization on that segment.

I find, however, that a definite plate is present on the larval skin.

: Flavithorax is probably the most recent derivative in the Flavicosta sub-

group, having arisen from flavicosta or its ancestor. It seems most likely that

the species arose in geographical isolation.

A specimen which is possibly a hybrid between flavicosta and flavithorax

is discussed in the systematics section of flavicosta. _

DISTRIBUTION. Coastal southwestern India and Ceylon. Material Exam-

ined: 31 specimens; 60°, 69, 2 P, 17 L; 2 individual rearings (1 larval, 1 in-

complete).

INDIA. Mysore: Karwar, Aug-Sept 1921, P.J. Barraud, © lectotype oe

1 1p (881), 1 1p (872), 4c, 5 ee one [BMNH].

No locality or date: 1 of [USNM ].

Additional records from the literature. CEYLON. Madola, near Opana-

yaka, Feb 1933, Henry (Barraud, 1934:103-104).

18. Orthopodomyia madrensis Baisas

Figs. 27, 28

1946. Orthopodomyia madrensis Baisas, 1946:41-45. TYPE: holotype * with

associated larval and pupal skins (THIB-4), Llavac, Laguna, Luzon, Phil-

ippine Islands, larva found in a moderately large treehole, Sept 1940, P.

Sunico [Location unknown].

Gneoabaa madrensis of Knight and Mattingly (1950:2, 17-20); Stone, Knight

and Starcke (1959:123); Delfinado, Viado and Coronel (1962: 438); Delfinado

(1966:68-70).

FEMALE. Wing: 3.44 mm. Proboscis: 2.17mm. Forefemur: 2.41 mm.

Abdomen: about 3.4 mm. Head: integument of head capsule, clypeus and ends

of most flagellar segments brown to dark brown, torus, basal flagellar seg-

ments and central portion of most remaining flagellar segments lighter; frontal

and orbital bristles brown; decumbent scales white except for yellowish ones in

front of dorsolateral broad scales; erect scales not longer and narrower ante-

riorly, posterior ones all brown, anterior ones white-tipped; dorsolateral

scales all yellow; lateral and ventral scales yellow or white tinged with yellow;

Zavortink: Genus Orthopodomyia 123

labium brown-scaled with cream-colored to yellow scales in narrow to broad

ring near middle, dorsal patch or complete ring near apex, ventral streak usu-

ally extending from or basad of middle ring to or near apex and sometimes in

streak on dorsal surface extending basad of middle ring; palpus 0.42 of probos-

cis, 9-segmented, segment 4 short, segment 5 minute, largely brown-scaled

with cream-colored or yellow scales over joint between segments 2-3 and some-

times near base of segment 2, and with white scales at apex of segment 3 and

covering all of segment 4 and 5; torus scales white or yellow-tinged; flagellar —

scales white. Thorax: mesonotum brown, greyish-brown or reddish-brown,

scutellum usually lighter and greyer, postnotum dark mesally, lighter lateral-

ly, pleuron uniformly brown, greyish-brown or reddish-brown, or with lighter

areas; paratergite bare; acrostichal scales brown or white at caudal end, re-

mainder mostly white but with some brown and yellow; dorsocentral scales

white at caudal end, remainder mostly brown but with some yellow; lateral pre-

scutellar scales all brown except for white near scutellum; posterior fossal and

supraalar scales white; fossal scales mostly brown but with some yellow; pra

with 1 bristle; mep bristles white; ppn scales all or mostly broad and flat, up-

per ones yellowish, remainder white; pra, ppl and stp scales white; pst scales

white or cream-colored; ssp scales in a long row, white; hypostigial scales

broad, flat and white and connected to ssp scale row; upper half of stp with 1

large conspicuous bare area opposite upper end of lower mep scale patch; lower

mep scales white. Legs: integument of mid- and hindcoxae same color as pleu-

ron or lighter in upper portion, forecoxa much lighter; coxal scales white ex-

.cept for some brown ones on lower portion of hindcoxa; forefemur with an ir-

regular mostly brown-scaled ventrally incomplete preapical ring, anterior sur-

face white- to yellowish-scaled at base, remainder largely brown-scaled with

white to yellowish speckles, posterior surface similar but with more light

scales and with light-scaled patch basad of incomplete preapical ring; mid- and

hindfemora with an irregular largely brown-scaled ventrally incomplete pre-

apical ring preceded basally by an irregular ring of mostly white or cream-col-

ored scales, anterior surface brown-scaled at base, remainder brown-scaled

with white to yellowish speckles, posterior surface usually mostly brown-scaled

basally, ventrally and dorsally basad of light ring, remainder largely pale-

scaled or sometimes posterior surface entirely pale-scaled except for some |

brown scales dorsally; tibiae largely brown-scaled with cream-colored spec-

kles and usually with an irregular narrow subbasal brown-scaled ring, posteri-

or surface of foretibia, posterior and sometimes part of dorsal surface of mid-

tibia and basal portion of posterior surface of hindtibia with largely cream-col-

ored scales, and apical portion of posterior surface of hindtibia brown-scaled;

tarsi largely brown-scaled, segment 1 with a few white speckles and sometimes

with ventral and/or posterior surface largely pale-scaled; foretarsal segment 3

and sometimes segment 2 white- to beige-scaled at ends and segments 4 and 5

all white- to beige-scaled; midtarsal segment 1 white- to beige-scaled at apex,

segments 2 and 3 white- to beige-scaled at ends and segments 4 and 5 all white-

to beige-scaled; hindtarsal segment 2 broadly white-ringe at apex and segments

3 and 4 entirely white-scaled or more commonly with a small to large brown

dorsal patch or complete ring distad of middle of segment 3. Wing: costal vein

with sectoral and accessory subcostal light patches, and usually with small dis-

tinct basal, prehumeral, humeral and presectoral light patches, or prehumer-

al, humeral and/or presectoral patches absent, or basal and prehumeral or

basal, prehumeral and humeral patches large and confluent; vein Sc without

presectoral light patch; vein R with or without basal light patch; vein Ry usually

124 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

with accessory subcostal light patch and rarely without subcostal light patch;

base of vein 1A dark; vein Cu without light patch basad of furcation; some or

all of veins Ro, Rg, R445, M1+2, M3+4 and Cuy with white scales at apex. Hal-

tere: integument white to brown; scales transparent to whitish. Abdomen: ter-

gite I with brown-scaled fan and scattered white scales laterally; remaining

segments with dark scales brown or black, light scales white or rarely yellow-

ish; tergites II-VII dark-scaled with basolateral and apicolateral light patches

which become smaller on distal segments, basolateral patches of same segment

sometimes connected by weak narrow light-scaled basal band, basal and apical

patches of same segment sometimes broadly connected along lateral edge on

proximal segments and narrowly connected on distal segments by light scales;

all or only more distal of tergites IV-VII with very small to moderately large

subdorsal median light patch; some tergites sometimes with middorsal

basal light patch; tergite VIII dark-scaled with narrow basal light band; ster-

nites IJ or II and III light-scaled; sternites III-V or IV and V light-scaled with

median lateral dark-scaled patch; sternites VI-VIII dark-scaled with light scales

in basolateral patch and sometimes in median streak.

MALE. Similar to female. Proboscis: with narrow ring of white to cream-

colored scales distad of level of distal end of palpal segment 2, with broad pre-

apical ring of cream-colored scales, both rings usually joined by ventral light

streak, and with or without light speckles on dorsal surface. Palpus: about

0.77 of proboscis; segment 3 without light speckles. Antenna: integument of

torus lighter laterally; flagellar segments 1-4 with tufts of long white scales.

Abdomen: sternite VIII white-scaled basally, laterally and sometimes apically;

Sidepiece with light and dark scales. :

MALE GENITALIA (fig. 27). Segment VIII: tergite lobe longer than broad,

Sides diverging apically, apex emarginate and with irregular small teeth. Seg-

ment IX: tergite with 0 or 1 bristle on each side. Sidepiece: basal mesal lobe

with 7 or 8 bristles which are not sharply divided into stout and fine ones; each

bristle not or only slightly longer than the 1 dorsad of it; mesal surface mem-

branous distad of basal mesal lobe; patch of strong sinuous specialized mesal

bristles present, specialized tergomesal bristles absent. Clasper: not strong-

ly curved in distal 0.25. Aedeagus: weakly sclerotized; somewhat ''angular-

ovate" in outline, sternal surface flared laterally on each side near apex; with

pair of nearly contiguous to widely separated teeth along edges of sternal sur-

face about 0.55-0.67 from base; without basal projection between parameres.

Proctiger: paraproct sclerotization with 3-5 apical teeth; cercal setae 3 or 4.

PUPA (fig.27). Abdomen: 3.23 mm. Trumpet: 0.50 mm. Paddle: 0.66

mm. Cephalothorax: mostly straw-colored, posterior portion of mesonotum

and upper part of wing case darker, tan; integumentary sculpturing weakly de-

veloped, reticulate; hair 1-C moderately developed, usually single or double;

2-C moderately developed, usually 2, 3b (2-4); 3-C strongly to very strongly

developed, usually 4-6b (3-8); 4,5-C very strongly developed, usually 3, 4b (2-

4); 6-C cephalad of 7-C; 7-C very strongly developed, usually double (1-3b);

8-C moderately to strongly developed, single; 9-C moderately to strongly de-

veloped, usually double or single (1-3b). Trumpet: dark brown except at ex-

treme base and apex; widened gradually from base; pinna large. Metanotum:

evenly light brown in color or with lateral part of haltere case lighter; integu-

mentary sculpturing weakly developed, reticulate; hair 10-C moderately to

strongly developed, usually 4-6b (3-7); 11-C moderately developed, usually

single, sometimes with fine branches near apex; 12-C strongly developed, usu-

ally 4, 5b (2-5); fourth pair of hairs never developed. Abdomen: anterior seg-

Zavortink: Genus Orthopodomyia 125

ments light brown, posterior segments tan; anterior portion and posterior edge

of anterior segments darker; integumentary sculpturing moderately developed

on anterior segments; hair I-II usually 9-13b; 2-II slightly thickened and mesad

of 5-II; 5-II long and single; 2-III, IV fine; 5-IV usually 3b (2, 3b); 5-V usually

double (single or double); 5-VI single; 9-VII 10-12b and with longest branch ex-

tending to or nearly to alveolus of 9-VIII; 9-VIII usually 16-18b (12-19) and with

longest branch reaching 0.6-0.7 distance to apex of paddle. Paddle: very light

straw-colored; longer than broad, external portion Somewhat angular; external

buttress straight in middle portion and with small marginal spicules; midrib

convex mesally or straight. :

LARVA (fig. 28). Head: 1.03 mm. Siphon: 1.16 mm. Anal Segment: 0.36

mm. Head: integumentary sculpturing absent; dark, tan to brown, posteriorly, |

lighter, straw-colored to tan, anteriorly and around imaginal eye, collar brown;

mental plate tan to brown, usually with 8-10 teeth on each side; 13-C moderate-

ly developed, usually 3-5b. Antenna: short, largely light brown to brown, light-

er at apex. Thorax: epidermal pigment strongly developed, purple; spicules

absent; 8-P usually 4-6b; 1-M,T single. Abdomen: segment VII with large

plate; hair 1-I short, usually double (2, 3b); 1-II short, usually single (single,

double); 6-I, II with individual branches coarse to near apex and barbed only in

basal portion; 1-III moderately long, single; 1-IV and 13-III-V moderately long,

double, 1 branch longer than other; 1-V long, single. Segment VIII: sclerotized

plate large but not ringing segment; anterior comb scales usually 23-27 (20-28),

posterior comb scales usually 8-10 (8-13); anterior comb scales fringed only

or with several apical members of fringe lengthened and spine-like; posterior

comb scales with 9-7 large apical spines bordered on each side by numerous

lesser spines which become smaller and fringe-like proximally; posterior row

of comb scales 0.78-0.97 length of anterior row; hair 3-VIII usually 8-11b (7-

11). Siphon: index usually 3.8-6.6; base of siphon brown to black, apex straw-

colored, remainder evenly brown to dark brown or lighter basad of alveolus of

hair 1-S; imbricate integumentary sculpturing weakly to moderately developed

basally; hair 1-S located 0.44-0.53 from base of siphon, usually 8-11b (6-13).

Anal Segment: largely light to dark brown in color with base and dorsal surface

darker; weakly to moderately developed imbricate integumentary sculpturing

present; dorsal anal gills about 1. 8-2.3 times length of ventral and 0. 8-1.1

length of anal saddle.

BIONOMICS. The immature stages have been collected in treeholes and in

a hole in the stump of a treefern. The immature stages of anopheloides may

occur with those of this species.

SYSTEMATICS. Adults of this species and the 6 following differ from flavi-

costa and flavithorax by the following features: (1) light scales on the femora,

tibiae and first tarsal segments scattered rather than in distinct patches, (2)

presence of accessory subcostal light patch on the costal vein and vein Rj, (3)

absence of a light patch basad of the fork on vein Cu in either sex, (4) absence

(usually) of a broad white ring over the joint between hindtarsal segments 1 and

2, (5) absence of scales on the paratergite, (6) scales on first 4 or 5 flagellar

segments in the male, and (7) the male palpus definitely shorter than the pro-

boscis.

The adults of madrensis are unique in the Albipes group in the presence of

a patch of hypostigial scales and share a bristle on pra with only flavicosta and

flavithorax. The white-scaling on the thorax of madrensis is usually a more

pure white than in the other species. Madrensis is not distinct from the other

members of the Albipes group in ornamentation of the proboscis and base of the

126 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

costa as stated by Baisas (1946:41) and Knight and Mattingly (1950:6,19). The

light-scaling of the proboscis is: more frequently white than yellow and is sub-

ject to the same variation in extent as in the other species; the light ornamen-

tation of the base of the costa is actually more variable in this species than in

any other, ranging from only a small basal patch to small and distinct basal,

prehumeral, humeral and presectoral patches or to large and confluent basal,

prehumeral and humeral patches. Only sternites II and sometimes III are en-

tirely white-scaled and in this respect madrensis differs only slightly from

other species. As is the case in albipes, andamanensis, anopheloides and si-

amensis, the number of all white hindtarsal segments varies. In madrensis

segments 3,4 and 5 may be all white or segment 3 may have a small dark patch

ora éompleté dark ring distad of the middle.

The male genitalia of this species are unique in the shape of the aedeagus

and in having all the bristles on the basal mesal lobe strong. They resemble

albipes in the development of a large patch of specialized mesal bristles.

The abdominal chaetotaxy of the pupa of madrensis is similar to flavicosta,

but the cephalothoracic chaetotaxy is similar to the Anopheloides subgroup.

The larva is more similar to siamensis than any other species. It can be told

from all other species of Orthopodomyia by the development of hairs 6-I, II on

the abdomen.

DISTRIBUTION. Endemic to Luzon. Material Examined: 56 specimens;

90°, 109, 8 P, 29 L; 2 individual rearings (1 larval, 1 incomplete).

REPUBLIC OF THE PHILIPPINES. Camarines Sur: Mt. Isarog, 20 Sept

1964, M. Delfinado, 1, 32, 6 P, 20 L[USNM], 60, 6 2 [BISHOP].

Laguna: Llavac, Sept 1940, P. Sunico (THIB), 1 [BISHOP]. Mt. Makiling,

8 June 1964 (IH 35), 3 L [UCLA]; 27 Aug 1966, 1 lpo (252-9) [UCLA].

Mountain: menice 6 Sept 1945 (455), 1 L TUSNM].

Quezon: Mt. Banahao, 14 Feb 1945 (P 624), 10, 2 L [USNM].

NO DATA. 1 1p (P 599-1) [USNM]; (P 605), 1 a [USNM].

Additional records from the literature. None.

19. Orthopodomyia siamensis, n. sp.

Figs. 29, 30

TYPES: Holotype: o with associated larval and pupal skins (TG 102-30),

Muang, Trang, Thailand, larva found in a roothole, 1964 [USNM]. Allotype: ¢

with associated larval and pupal skins (TG 111-31), Muang, Trang, Thailand,

larva found in a treehole, 1964 [USNM]. Paratypes: 2 po (TG 111-101, 102),

3 p? (TG 111-100, 103, 104), 1% (TG 111), 1 L (TG 111), same data as allotype.

FEMALE. Wing: 3.50 mm. Proboscis: 2.03 mm. Forefemur: 2.14 mm.

Abdomen: about 2.5mm. Head: integument of head capsule, clypeus and ends

of most flagellar segments brown to dark brown, torus lighter, middle portion

of most flagellar segments much lighter; frontal and orbital bristles brown; de-

cumbent scales all white or white anteriorly and yellow posteriorly and lateral-

ly; erect scales not longer and narrower anteriorly, brown posteriorly, white

or cream-colored or tipped anteriorly; dorsolateral scales brown anteriorly,

white or yellow posteriorly; lateral and ventral scales white tinged with yellow;

labium mostly brown-scaled, cream-colored scales in ring distad of middle, in

ventral streak from or basad of ring to near apex, and in dorsal patch near

apex; palpus 0.42 of proboscis, 4-segmented, segment 4 short, mostly brown-

Zavortink: Genus Orthopodomyia | 127

scaled, white scales over joint between segments 2-3, at apex of segment 3 and

covering all of segment 4; torus scales white to yellow-tinged; flagellar scales

white. Thorax: integument of mesonotum and postnotum light brown to brown

or greyish- or reddish-brown, scutellum often lighter and greyer, pleuron

darker; paratergite bare; acrostichal scales brown at caudal end, remainder

brown and yellow; dorsocentral scales yellow or yellow and white at caudal end,

remainder yellow and brown; lateral prescutellar scales brown except for white

laterally and near scutellum; posterior fossal scales white and yellow; fossal

scales mostly brown anteriorly, yellow posteriorly; supraalar scales white

mesally and golden laterally; pra bristles absent; mep bristles white; ppn with

upper scales narrow and curved, at least a few lower ones broad, yellow or

yellow and white; pra with cream-colored scales; ppl, pst, ssp and stp scales

white to cream-colored or yellow-tinged; ssp scales in a short to moderately

long row; hypostigial scales absent; upper half of stp with bare areas opposite

upper and middle portions of lower mep scale patch; lower mep scales white.

Legs: integument of mid- and hindcoxae same color as pleuron, forecoxa light-

er; forecoxal scales white or some yellow-tinged, mid- and hindcoxal scales

white above, brown below; forefemur with an irregular largely brown-scaled

preapical ring, anterior surface with cream-colored to yellow scales at base,

remainder brown-scaled with cream-colored to yellow speckles, posterior sur-

face brown-scaled at base and with cream-colored to yellow scales in a patch

basad of preapical ring, remainder brown-scaled with cream-colored to yellow

speckles; midfemur with an irregular largely brown-scaled complete or incom-

plete preapical ring usually preceded basally by an irregular dorsally incom-

plete ring of mostly white to cream-colored scales, anterior surface brown-

scaled at base and ventrally to beyond middle, remainder brown-scaled with

cream-colored to yellow speckles, posterior surface brown-scaled basally and

ventrally to beyond middle, brown-scaled with cream-colored to yellowish

speckles dorsally, remainder largely pale yellowish-scaled; hindfemur with

brown and cream-colored rings similar to those of midfemur, anterior surface

largely brown-scaled, having only a few cream-colored to yellowish speckles,

posterior surface like that of midfemur or nearly entirely brown-scaled; tibiae

largely brown-scaled with cream-colored to yellow speckles, white to cream-

colored scales at base, brown scales in a narrow subbasal ring, posterior sur-

face of fore- and midtibiae and basal portion of posterior surface of hindtibia

with mostly cream-colored scales, apical portion of posterior surface of hind-

tibia largely brown-scaled, and anterior surface of hindtibia cream-colored

with brown speckles; tarsi largely brown- to black-scaled, segment 1 with a

few cream-colored to yellow speckles and sometimes with ventral and/or pos-

terior surface largely pale-scaled; fore- and midtarsal segments 2 and 3 white-

to beige-scaled at ends and segments 4 and 5 entirely white- to beige-scaled;

mid- and hindtarsal segment 1 white-scaled at apex; hindtarsal segment 2

broadly white-ringed at apex and segments 3 and 4 entirely white-scaled or less

commonly with dark-scaled ring distad of middle of segment 3. Wing: costal

vein with sectoral and accessory subcostal light patches, and usually with small

distinct basal, prehumeral, humeral and presectoral light patches, or basal and/

_or preSectoral patches absent, or basal, prehumeral and humeral patches large

and confluent; vein Sc without presectoral light patch; vein R with or without

basal light patch; vein Ry with accessory subcostal light patch; vein 1A dark at

base; vein Cu without light patch basad of furcation; some or all of veins Ro,

R3, R4+5, Myj+2, M3+4, and Cuj with white scales at apex. Haltere: integu-

ment of stem whitish to tan, knob beige to brown; scales transparent to whitish

128 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

and brownish. Abdomen: tergite I with brown-scaled fan and with scattered

white scales; remaining segments with dark scales brown to black, light scales

white or white and yellow; tergites II-VII largely dark-scaled with basolateral

and apicolateral light-scaled patches, lateral patches of same segment connected

along lateral edge on proximal segments, basal patches of same segment some-

times connected by basal light band on distal segments, and with median sub-

dorsal light-scaled patch on all or more distal tergites; tergite VIII with light-

scaled basal band; sternites II-VIII dark-scaled with basolateral light-scaled

patch which becomes smaller on distal segments.

MALE. Similar to female. Proboscis: white scales in complete narrow

ring distad of distal end of palpal segment 2 and in dorsal preapical patch con-

nected laterally to 1 or 2 preapical ventral patches, or with broad yellowish

preapical ring which extends further basally on ventral surface than on dorsal

surface. Palpus: about 0.82 of proboscis; segment 3 not light-speckled. An-

tenna: integument of torus lighter laterally than in female; flagellar segments

1-4 with tufts of long white scales. Abdomen: sternite VIII light-scaled basally

and apically; sidepiece with light and dark scales.

MALE GENITALIA (fig. 29). Segment VIII: tergite lobe about as long as

broad to longer than broad, usually parallel sided, apex rounded or emarginate,

with or without irregular teeth. Segment IX: tergite with 0 or 1 bristle on each

side. Sidepiece: basal mesal lobe with 4 or 5 stout bristles and 1 or 2 finer;

each stout bristle only slightly longer than the one dorsad of it; mesal surface

sclerotized distad of basal mesal lobe; specialized mesal bristles absent, 1 or

2 specialized tergomesal bristles present. Clasper: not strongly curved in dis-

tal 0.25. Aedeagus: strongly sclerotized; broadest about 0.55-0.60 from base,

tapering to obtuse apex, sides sometimes almost parallel to near base, basal

portion truncate; I pair of very strongly sclerotized, nearly contiguous sternal

teeth about 0.50-0.55 distance from base; without basal projection between

parameres. Proctiger: paraproct sclerotization with 3 or 4 apical teeth; cer-

cal setae 3 or 4.

PUPA (fig.29). Abdomen: 2.99 mm. Trumpet: 0.46 mm. Paddle: 0.62

mm. Cephalothorax: mostly light straw-colored, posterior portion of mesono-

tum and posterior portion of wing case darker, tan; integumentary sculpturing

absent; hairs 1, 2-C strongly developed, usually 6, 7b (5-8); 3-C strongly de-

veloped, usually 7-9b (4-10); 4,5-C strongly developed, usually 5, 6b (3-8); 6-

C cephalad of 7-C; 7-C strongly developed, double; 8-C strongly developed,

usually 6, 7b (3-8); 9-C strongly developed, 3-5b. Trumpet: light brown in

color with apex and base lighter; quite similar to wilsoni in shape, narrower

than in most other species, parallel sided or widening slightly towards apex,

pinna small. Metanotum: evenly tan; integumentary sculpturing absent; hair

10-C moderately developed, usually 7-10b (7-13); 11-C moderately developed,

usually single; 12-C strongly developed, 6-9b; fourth pair of hairs never devel-

oped. Abdomen: anterior segments tan, posterior segments straw-colored;

anterior portion and posterior edge of anterior segments darker; integumen-

tary sculpturing moderately developed on anterior segments; hair 1-II 11-19b;

2-II quite thickened and laterad of 5-II; 5-II long and single; 2-H, IV thickened;

5-IV double; 5-V, VI single; 9-VII 9-12b and with longest branch extending to

alveolus of 9-VIII; 9-VIII 13-16b and with longest branch reaching 0.8-0.9 dis-

tance to apex of paddle. Paddle: very light straw-colored; longer than broad,

external portion sometimes Somewhat angular; external buttress nearly straight

in middle portion and with marginal spicules; midrib straight to slightly convex

mesally.

Zavortink: Genus Orthopodomyia 129

LARVA (fig. 30). Head: 0.97 mm. Siphon: 1.27 mm. Anal Segment: 0.37

mm. Head: integumentary sculpturing absent; largely tan to light brown in col-

or, darker posteriorly, lighter around imaginal eye, collar dark brown to black;

mental plate tan to light brown, usually with 8-10 teeth on each side; 13-C mod-

erately strong, usually with 8, 9b (8-11). Antenna: moderately long, brown

basally, lighter apically. Thorax: epidermal pigment strongly developed, pur-

ple; spicules absent; 8-P usually 2,3b; 1-M,T single. Abdomen: segment VI

with dorsal sclerotized plate and segment VII with large sclerotized plate; hairs

1-I, II moderately long, single; 6-I, IJ with individual branches tapering to fine

apex and barbed from near base to near apex; 1-III long, single; 13-III-V mod-

erately long, usually double (double, single), and with 1 branch longer than

other; 1-IV long, double; 1-V very long, double. Segment VIII: sclerotized

plate large but not ringing segment; anterior comb scales usually 20-27, pos-

terior comb scales usually 8-10 (7-11); larger anterior comb scales shaped like |

posterior comb scales, with 5 or 6 large apical spines bordered on each side by

numerous-lesser spines which become smaller and fringe-like proximally; pos-

terior row of comb scales about 0. 80-0. 94 length of anterior row; hair 3-VIII

usually 8-10b. Siphon: index usually 6.4-7.8 (6.4-10.1); base of siphon brown

to dark brown, apex straw-colored, remainder uniformly light to deep brown;

weakly developed imbricate integumentary sculpturing present basally; hair 1-S

located 0.41-0.49 from base of siphon, usually 6-8b (5-8). Anal Segment: sad-

dle brown to dark brown with base and dorsal surface darker; weakly to mod-

erately developed imbricate integumentary sculpturing present; dorsal anal

gills 1.2-2.0 length of ventral and about 1.0-1.6 times length of anal saddle.

BIONOMICS. The immature stages have been collected in rot holes in

trees, stumps and roots. So far as is known, they have not been collected in

association with any other species of the Albipes group, although the treehole-

breeding andamanensis and anopheloides occur in the same area.

SYSTEMATICS. On larval and pupal characters this species definitely be-

longs to the Flavicosta subgroup, but the adults cannot be adequately separated

from those of the Anopheloides subgroup and the male genitalia arevery similar

to those of papuensis.

Adults of this species show the same hindtarsal markings as thadrennia

and the range in costal ornamentation is almost as great as in that species. In

published keys to species of this group (Knight and Mattingly, 1950:6), adults of

Siamensis would run to either andamanensis or maculipes.

The larva resembles madrensis and the pupa flavithorax. The larva of si-

amensis is apparently the only one in the Albipes group to develop a sclerotized

plate on abdominal segment VI. |

DISTRIBUTION. Peninsular Thailand. Material Examined: 62 specimens;

l1o%, 69, 15P, 30L; 16 individual rearings (7 larval, 8 pupal, 1 incomplete).

THAILAND. Narathiwat: Khau Lau (00430), 1 pc" (00430- 100), 1L [USNM].

Trang: Muang (TG 20), 1 1p? (20-52), 1 p? (20-100) [USNM]; same local-

ity (TG 98), 1 10 (98-47) [USNM]; same locality (TG 102), lpo’ holotype (102-

30) [USNM]; same locality (TG111), 1p? allotype (111-31), 2 po’ (111-101, 102),

3 p¢ (111-100, 103, 104), 1%, 1 L [USNM].

No locality or date: (00476), 7 L [USNM]; (00477), 3 L [USNM]; (00481),

4L [USNM]; (NV 12), 1 lpo (12-13), 1L [USNM]; (NV 13), 1 1po (13-10), 4 L

[USNM]; (NV 21), 1 lpo (21-11), ‘po (21-112) [USNM]; (RN 49), 1 lpo (49-

1-10) [USNM]; (TG 121), 1 L [USNM].

Additional records from the literature. None.

130 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

WILSONI SUBGROUP

20. Orthopodomyia wilsoni Macdonald

Figs. ob; 32

1958. eet hans wilsoni Macdonald, 1958:121-124. *TYPE: holotype &

with associated larval and pupal skins (0348/4), 15 to 16 miles north of

Kuala Lumpur in the Ulu Gombak Forest Reserve, Selangor, Malaya,

larva found in a bamboo internode, 8 Oct 1957 [BMNH].

Orthopodomyia wilsoni of Stone, Knight and Starcke (1959:124); Macdonald and

Traub (1960:94).

FEMALE. Wing: 2.84 mm. Proboscis: 1.65 mm. Forefemur: 1.68 mm.

Abdomen: about 2.1 mm. Head: integument of head capsule, clypeus, torus

and ends of most flagellar segments brown to dark brown, middle portion of

most flagellar segments lighter; frontal and orbital bristles brown; decumbent

scales white to cream-colored anteriorly, yellowish posteriorly; erect scales

not longer and narrower anteriorly, brown posteriorly, white anteriorly; dor-

solateral scales brown anteriorly, yellowish posteriorly; lateral and ventral

scales yellowish; labium largely brown-scaled with cream-colored scales in

ring at about 0.75 distance from base, in ventral streak extending slightly bas-

ad and distad of ring and in dorsal patch between ring and apex; palpus 0. 42 of

proboscis, 4-segmented, segment 4 short, largely brown-scaled with white

scales over joint between segments 2-3 and covering segment 4; torus scales

white to yellow-tinged; flagellar scales white. Thorax: integument of mesono-

tum and pleuron brown, scutellum and sometimes postnotum lighter; parater-

gite bare; acrostichal scales brown at caudal end, remainder mostly yellow but

with some brown and sometimes white; dorsocentral scales white and yellow at

caudal end, remainder brown and yellow; lateral prescutellar scales brown or

brown and yellow except for white scales laterally and posteriorly; posterior

fossal scales yellowish to yellow; fossal scales brown and yellow; supraalar

scales white mesally, golden laterally; pra bristles absent; mep bristles white;

ppn with yellowish to golden scales, some lower ones broad, flat; pra, ppl, pst,

ssp, stp and lower mep scales cream-colored to yellow; ssp scale row usually

short, sometimes moderately long; hypostigial scales absent; upper half of stp

with bare areas opposite upper and middle portions of lower mep scale patch.

Legs: integument of mid- and hindcoxae same color as pleuron or slightly light-

er, forecoxa always lighter; forecoxal scales white to yellowish, midcoxal

scales white to yellowish above and brown below, hindcoxal scales white above

and golden or brown below; femora similar to those of siamensis, but midfe-

mur sometimes not brown-scaled to beyond middle on ventral anterior surface,

hindfemur often with numerous light speckles on anterior surface and sometimes

not brown-scaled at base of posterior surface; tibiae similar to those of siamen-

sis, but mid- and hindtibiae with a more or less conspicuous ring of white to

cream-colored scales distad of subbasal brown-scaled ring and hindtibial pos-

terior surface sometimes all light-scaled; tarsi largely brown- to black-scaled,

segment 1 with cream-colored speckles at least on hindleg and sometimes on

all three legs and at least hindtarsal segment 1 sometimes with ventral surface

largely pale-scaled; foretarsal segments 2-5 all dark-scaled, or segments 2

and 3 white-scaled at base and segments 4 and 5 with brown scales lighter than

more proximal segments; midtarsal segments 2 and 3 white-scaled basally and

Zavortink: Genus Orthopodomyia 131

_ dark- or beige-scaled apically, segments 4 and 5 white- to beige-scaled; hind-

tarsal segment 1 usually dark-scaled at apex but sometimes with narrow white-

scaled ring, segments 2 and 3 with narrow to broad white-scaled ring at each

end, segment 4 entirely white-scaled. Wing: costal vein usually with sectoral

and accessory subcostal light patches, and usually with small distinct basal,

prehumeral, humeral and presectoral light patches, or humeral and presectoral

patches large and confluent; vein Sc without presectoral light patch; vein R

without basal light patch and rarely without sectoral light patch; vein Ry with

accessory subcostal light patch; vein 1A dark at base; vein Cu without light

patch basad of furcation; some or all of veins Ro, Rg, R4+5, M1i+2, M3+4 and

Cul with white scales at apex. Haltere: integument of stem whitish, knob beige

to tan; scales transparent to whitish. Abdomen: tergite I with brown-scaled

fan and without or with only a few scattered light or dark scales laterally; re-

maining segments with dark scales brown to blackish, light scales white, yel-

low or golden, more frequently yellow or golden than in most other species;

tergites II-VII dark-scaled with basolateral and apicolateral light-scaled patches,

apicolateral light patch usually quite yellow or golden, lateral patches of seg-

ment II or segments II and III broadly joined; tergite II or II and III with large

to very large basal middorsal light (usually yellowish) patch which may be con-

nected to basolateral patches by weak light-scaled basal band; tergites IV-VII

with scattered light scales basally or with basal light band which is more strong-

ly developed on distal segments; tergites III-VII with subdorsal median light

patch (usually yellow or golden); tergite VIII dark-scaled with light basal band;

sternites II-VII dark-scaled with basolateral light patch, patch large on segment

II, smaller on distal segments; sternite VIII scales dark. )

MALE. Similar to female. Proboscis: with narrow incomplete to complete

white- and yellow-scaled ring just distad of distal end of palpal segment 2, with

dorsal preapical yellow-scaled patch, and sometimes with yellow-scaled pre-

apical patch on ventral surface basad of dorsal patch, the 2 patches connected

laterally when ventral is present. Palpus: about 0.84 of proboscis; segment 3 |

not speckled. Antenna: integument of torus lighter laterally; flagellar segments

1-4 with tufts of long white scales. Abdomen: tergites more frequently and

more broadly banded and subdorsal median light-scaled patches larger; sternite

VIII with light scales apically and basally; sidepiece with light and dark scales.

MALE GENITALIA (fig. 31). Segment VII: tergite lobe longer than broad,

parallel sided or broader apically, apex truncate or emarginate and with small

teeth. Segment IX: tergite with 0 or I bristle on each side. Sidepiece: basal

mesal lobe with 4 stout bristles and 1 finer; each stout bristle considerably ©

longer than the 1 dorsad of it; mesal surface weakly sclerotized distad of basal

mesal lobe; specialized mesal bristles absent; sometimes 1 or 2 specialized

tergomesal bristles present. Clasper: not strongly curved in distal 0.25. Ae-

deagus: moderately sclerotized; pyriform, with 1 pair of strong, sternal teeth

near apex, teeth always separated; without basal projection between parameres.

Proctiger: paraproct sclerotization with 3 or 4 apical teeth; cercal setae 2-4.

PUPA (fig.31). Abdomen: 3.19mm. Trumpet: 0.56 mm. Paddle: 0.64

mm. Cephalothorax: evenly yellowish-tan colored; integumentary sculpturing

absent; hairs 1-3-C moderately developed, usually double (1-3b); 4-C moder-

ately developed, usually single (1-4b); 5-C moderately developed, usually sin-

gle or double (1-3b); 6-C cephalad of 7-C; 7-C only hair longer than trumpet,

very strongly developed, usually double (2, 3b); 8,9-C moderately developed,

usually single (single, double). Trumpet: dark brown in color, base and apex

lighter; narrow and parallel sided throughout most of its length; pinna small.

132 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

Metanotum: evenly yellowish-tan colored or with central portion slightly darker

and haltere case slightly lighter; integumentary sculpturing absent; hair 10-C

moderately developed, usually 5, 6b (5-9); 11-C moderately developed, usually

3, 4f (1-5); 12-C moderately developed, usually single, sometimes with a couple

of fine branches beyond middle; fourth pair of hairs never developed. Abdomen:

all segments yellowish-tan colored; anterior portion and posterior edge of seg-

ments II and IV sometimes slightly darker; integumentary sculpturing weakly

developed on anterior segments; hair 1-II moderately developed, dendritic or

with 1 or more stalks from which 18-34 fine ultimate branches arise; 2-II fine

and mesad of 5-II; 5-II long, single; 2-III,IV fine; 5-IV usually 3, 4b; 5-V usu-

ally 2, 3b and extending to or almost to alveolus of 4-VII; 5-VI usually double

(1-3b) and extending to alveolus of 4-VIII; 9-VII usually 8-10b and with longest

branches extending to or nearly to alveolus of 9-VIII; 9-VIII 11-14b and with

longest branch reaching about 0.75 distance to apex of paddle. Terminal Seg-

ments: male genital lobe projecting only slightly beyond median caudal lobe.

Paddle: light straw-colored; longer than broad, narrowly elliptical or oval in

outline; external buttress nearly straight to slightly convex laterally in distal

portion and with small marginal spicules; midrib slightly convex mesally.

LARVA (fig.32). Head: 0.97 mm. Siphon: 1.81 mm. Anal Segment: 0.33

mm. Head: distinctly broader than long; integumentary sculpturing absent;

largely straw-colored to tan, lighter around imaginal eye, darker posteriorly,

collar dark brown; mental plate tan, usually with 8,9 teeth on each side; hairs

4-10-C not all equally developed; 13-C moderately strong, usually with 6, Tb

(5-8). Antenna: moderately long, evenly brown to dark brown or lighter apical-_

ly. Thorax: epidermal pigmentation unknown; spicules absent; hairs 0,1,3-7-P

not all equally developed; 8-P usually 5-7b (5-9); 1-M moderately long, usual-

ly 3, 4b (1-5); 1-T moderately long, usually 2,3b (1-5); 13-T moderately strong,

usually 5, 6b (4-7). Abdomen: segment VII with large sclerotized plate; hair

1-I, II moderately long, usually 3, 4b (1-4); 6-I, II with individual branches finer

than 6-III, tapering to fine apex and barbed from near base to near apex; I-III

moderately long, usually 2, 3b (1-3); 13-III moderately long, usually 3, 4b (2-4);

1-IV moderately long, usually 3, 4b (1-5); 13-IV, V moderately long, usually 5,

6b (2-7); 1-V very long, single. Segment VIII: sclerotized plate large but not

ringing segment; anterior comb scales usually 21-24 (20-26), posterior comb

scales usually 10-12 (7-14); larger anterior comb scales shaped like posterior

comb scales, with 1 long apical spine and a fringe, the distal elements of the

fringe enlarged, about 0.20-0.50 as long as terminal spine; posterior row of

comb scales about 0.71-0.99 length of anterior row; hair 3-VIII usually 8-10b

(6-11). Siphon: index usually 7.2-11.5; base of siphon dark brown to black,

portion between base and alveolus of hair 1-S light brown to tan, portion be-

tween 1-S and apex darker, brown to dark brown, extreme apex not or slightly

lighter; weakly developed imbricate integumentary sculpturing present basally;

hair 1-S located 0.30-0.37 from base of siphon, usually 8-10b (6-10). Anal Seg-

ment: largely light brown to brown in color, dorsal surface and base sometimes

darker; integumentary sculpturing absent or very weakly developed and imbri-

cate; dorsal anal gills 1.4-2.2 times length of ventral and usually 1.1-1.6 (1.1-

3.0) length of anal saddle.

BIONOMICS. Larvae have been collected in bamboo internodes with mod-

erate to large holes or cracks, but have not been collected in bamboo stumps.

Although wilsoni occurs in the same habitat as albipes in Malaya, the two spe-

cies have never been found in the same collection. Larvae were once collected

in the fungus Polystictus xanthopus. Nothing is known of the habits of the adults.

Zavortink: Genus Orthopodomyia _ 133

SYSTEMATICS. Adults are distinct in hindtarsal banding and in the pres-

ence of a large, usually yellowish, middorsal basal patch on abdominal tergite

II and sometimes III. The males of other species may approach wilsoni in the

latter character, but the patch is usually smaller and not yellow. The abdomi- |

nal light-scaling is generally more yellow or golden on all segments in wilsoni

than in any other species, and the ssp scale line is sometimes reduced to only

3 or 4 scales.

The male genitalia are similar to albipes but are distinct in lacking spe-

cialized mesal bristles. The genital lobe of the male pupa is very small and

diagnostic in wilsoni.

The pupa is differentiated from others in the Albipes group by short and

weakly developed cephalothoracic setae (except for hair 7-C) and a cylindrical

darkly-pigmented trumpet. Its abdominal chaetotaxy is similar to albipes. The

most distinctive feature of the larva is the shape of the comb scales. The si-

phon is longer and thinner than in any other species except for occasional spec-

imens of Siamensis and papuensis. The modal condition of the larval chaeto-

taxy is rather distinctive, but some individuals have the number of branches in

hairs 1-M, T, I-IV and 13-T, II-V reduced, and hair 1 may even be single. The

modal chaetotaxy is rather similar to that of the Central American phyllozoa.

This appears to be a relictual species; it has no close relatives, but its

affinities probably lie with albipes. I believe this species is distinctive enough

to be placed in a separate subgroup.

DISTRIBUTION. Malay Peninsula. Material Examined: 45 specimens;

90°, 69, 13 P, 17 L; 13 larval individual rearings.

FEDERATION OF MALAYSIA. Selangor: Tembler Park, 25 June 1958

(0672), 1 lpo (0672-1) [LIVER]. Ulu Gombak, 21 Aug 1956, J. A. Reid (0548),

1 9 [LIVER]; 8 Oct 1957, W.W. Macdonald (0348), 1po holotype (0348-4), 2 1po

(0348-8, 14), 3 lp? (0348-1, 5,6) {[BMNH], 2 1lpo (0348-9, 15), 2 1p? (0348-11,

12) [USNM]; 15 Jan 1958 (0459), 2 1po (0459-1, 3) [LIVER].

THAILAND. No locality or date: (RN 7), 1L [USNM]; (RN 8), 3L [USNM];

(RN 36), 1% [USNM].

Additional records from the literature. None.

ALBIPES SUBGROUP

Ys Orthopodomyia albipes Leicester

Figs. 33, 34

1904. Orthopodomyia albipes Leicester, 1904:237-239. *TYPE: lectotype &

with genitalia slide, specimen labeled "Orthopodomyia albipes (type) Lei-

cester,'' Pahang Road, 5 3/4 miles from Kuala Lumpur, [Selangor], Ma-

laya, reared from a larva collected in a bamboo internode, 8 Apr 1903

(date of eclosion), PRESENT SELECTION [BMNH]. Stated to be proba-

bly only a variety of anopheloides by Edwards (1913:239); given specific

rank by Edwards (1926:117, 118); considered as a form of anopheloides

by Barraud (1927:527, 528, 529); resurrected by Edwards (1928:53).

Orthopodomyia albipes of Theobald (1907:527-530); Daniels (1908:4); Leicester

(1908:176-177, in part); Theobald (1910a:30; 1910b:470); Edwards (1913:239;

1922:458, 470; 1926:117, 118; 1928:53,57; 1932:108); Barraud (1934:102);

134 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

Baisas (1946:34, 35,45); Brug and Bonne-Wepster (1947:183); Knight and

Mattingly (1950:16-17); Macdonald (1957:14; 1958:123, 124, 125); Stone, Knight

and Starcke (1959:122); Macdonald and Traub (1960:94); Delfinado, Viado and

Coronel (1962:438); Delfinado (1966:68); Stone, Scanlon, Bailey, Definado

and Bram (1966:82).

Orthopodomyia anopheloides form albipes of Barraud (1927:527, 528, 529); Borel

(1930a:298-302; 1930b:167).

Finlaya albipes of Giles (1904:366).

FEMALE. Wing: 3.24mm. Proboscis: 2.05 mm. Forefemur: 2.22 mm.

Abdomen: about 2.5mm. Differs from siamensis as follows. Head: decum-

bent scales all white; posterior dorsolateral, lateral and ventral scales white;

labial markings as in siamensis, but pale scales white and ring at about middle

of proboscis; palpus 0.58 of proboscis, 5-segmented, segment 4 moderately

long, segment 5 short, largely brown-scaled with white scales over joints be-

tween segments 2-3, 3-4 and 4-5, covering all of segment 5 and sometimes

near joint between segments 1-2; torus scales white. Thorax: acrostichal

scales usually brown, sometimes yellow, at caudal end, remainder largely yel-

low and brown, but usually with some white; dorsocentral scales white or white

and yellow at caudal end, remainder yellow and/or cream-colored and brown;

lateral prescutellar scales sometimes yellow mesally; posterior fossal scales

white or white and yellow; supraalar scales white mesally and sometimes at

ends of anterior and middle projections, remainder yellowish, yellow or gold-

en; ppn with upper scales narrow and curved, lower ones narrow and curved or

broad and flat, upper yellow-tinged, lower white; pra, ppl, pst, ssp, stp and

mep scales white or sometimes cream-colored; ssp scales in a moderately

long line; stp with bare areas opposite upper and middle portions of lower mep

scale patch and usually also below ssp scales. Legs: forecoxal scales entirely

white to cream-colored; forefemur sometimes with small light-scaled patch

-basad of preapical brown-scaled ring on anterior surface and sometimes with

brown-scaling at base of posterior surface reduced; midfemur anterior surface

not always brown-scaled ventrally to beyond middle and posterior surface with

pale-scaling yellowish or cream-colored; mid- and hindtibiae sometimes with

more or less conspicuous largely white- to yellowish-scaled ring distad of sub-

basal brown-scaled ring, foretibia usually with an anterior light-scaled patch

in same position, anterior surface of hindtibia with either light or dark scales

predominating; tarsal segment 1 dark-scaled at apex; fore- and midtarsal seg-

ments 2 and 3 and sometimes 4 and 5 white-scaled at base, segments 4 and 9,

especially on midtarsus, sometimes light brown-scaled; hindtarsal segment 2

with small white-scaled dorsal patch or complete ring at base, apex brown-

scaled, with a few white scales or with complete narrow white ring, segments

3 and 4 entirely white-scaled, or rarely segment 3 with dorsal dark patch bas-

ad of middle. Wing: costal vein usually with accessory subcostal light patch,

and usually with small distinct basal, prehumeral, humeral and presectoral

light patches, rarely with prehumeral and humeral light patches absent; vein

Sc often with presectoral light patch; vein Ry usually with accessory subcostal

light patch; vein 1A white-scaled at base. Haltere: scales transparent to whit-

ish. Abdomen: tergite I fan sometimes with white scales; light scales usually

white, only infrequently some yellow; tergites II-V sometimes with a few light

scales basally; sternite II mostly light-scaled, sternites IHI-VII often with nar-

row basal light band, sternites II-VI or II-V with ventral apical light patch.

MALE. Similar to female; differs from siamensis as follows. Proboscis:

Zavortink: Genus Orthopodomyia : ; 133

always with complete broad white or rarely yellowish preapical ring which ex-

tends further basally along ventral surface than along dorsal surface. Palpus:

about 0.75 of proboscis. Abdomen: sternite VIII light-scaled basally, lateral-

ly and apically; sidepiece with all dark scales.

MALE GENITALIA (fig. 33). Segment VIII: tergite lobe always much longer

than broad and wider apically than basally, apex rounded or truncate and with

small serrations. Segment IX: tergite with 0-2 bristles on each side: Side-

piece: basal mesal lobe with 4 or 5 stout bristles and 1 or 2 finer; each stout

bristle slightly to considerably longer than the 1 dorsad of it; mesal surface

with or without long membranous area distad of basal mesal lobe; mesal sur-

face with patch of moderately to strongly developed sinuous or curved special-

ized bristles; specialized tergomesal bristles absent. Clasper: not strongly

curved in distal 0.25. Aedeagus: moderately sclerotized; pyriform, with1 pair

of strong sternal teeth near apex, teeth always separated; without basal projec-

tion between parameres. Proctiger: paraproct sclerotization with 3-5 apical

teeth; cercal. setae 2-4. 7

PUPA (fig. 33). Abdomen: 3.38 mm. Trumpet: 0.46 mm. Paddle: 0.62

mm. Cephalothorax: mostly light straw-colored, posterior portion of mesono-

tum and upper part of wing case slightly darker; integumentary sculpturing ab-

sent; hair 1-C moderately developed, usually 2-4b (2-5); 2-C strongly devel-

oped, usually 2, 3b (1-3); 3-C very strongly developed, usually 4-6b (4-9); 4-C

strongly developed, usually 3, 4b (2-4); 5-C very strongly developed, usually

4, 5b (3-6); 6-C cephalad of 7-C; 7-C very strongly developed, usually 3, 4b

(3-5); 8-C moderately developed, single or double; 9-C moderately developed,

usually single. Trumpet: largely brown to dark brown in color, base and apex

lighter; broadening rapidly from the base and attaining its maximum or nearly

maximum width in the basal 0.33, middle 0.34 almost or actually parallel sided;

pinna large, making up the distal 0.33. Metanotum: mostly light tan to tan in

color, haltere case lighter; integumentary sculpturing absent; hair 10-C mod-

erately developed, usually 2, 3b (2-5); 11-C moderately developed, usually 3, 4f

(2-5); 12-C moderately developed, usually 2, 3b (1-3); fourth pair of hairs never

developed. Abdomen: anterior segments straw-colored to tannish-brown, pos-

terior segments lighter, light straw-colored to yellowish-tan; anterior and

mesal portions and posterior edges of anterior segments darker, tan to light

brown; integumentary sculpturing moderately developed on anterior segments;

hair 1-II short, not extending to 0-III and usually with a central stalk from

which 19-35 very fine dendritic or forked branches arise; 2-II fine and mesad

of 5-II; 5-II long, single; 2-III, IV fine; 5-IV usually 3-5b; 5-V usually 3b (2-4)

and extending to alveolus of 4-VII; 5-VI usually 2b (2-5) and extending to alveo-

lus of 4-VIII; 9-VII usually 9-11b (8-13) and with longest branches reaching al-

veolus of 9-VIII; 9-VIII usually 12-14b (11-16) and with longest branch reaching

about 0.75 distance to apex of paddle. Terminal Segments: male genital lobe

projecting considerably beyond median caudal lobe... Paddle: very light straw-

colored; longer than broad, narrowly elliptical to obovate in outline; external

buttress nearly straight in distal portion and almost smooth; midrib slightly

convex mesally. |

LARVA (fig.34). Head: 1.15 mm. Siphon: 1.74 mm. Anal Segment: 0.43

mm. Head: distinctly broader than long; integumentary sculpturing absent;

largely straw-colored, lighter around imaginal eye, darker posteriorly, collar

brown; mental plate tan, usually with 11,12 (11-13) teeth on each side; hairs

4-10-C not all equally developed; 13-C moderately developed, usually 4-6b.

Antenna: moderately long, light brown in color. Thorax: epidermal pigment

136 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

strongly developed, purple; spicules absent; hairs 0,1,3-7-P not all equally

developed; 8-P usually 8-13b; 1-M long, usually 1-3b (1-4); 1-T long, usually

1-3b; 13-T strongly developed, usually 7-9b (5-10). Abdomen: segment VII

with large sclerotized plate; hairs 1-I, I moderately long, usually 2, 3b; 6-I, II

with individual branches finer than 6-III, tapering to fine apex and barbed from

near base to near apex; 1-III long, usually 2, 3b (1-4); 13-III-V moderately long,

usually 4, 5b (3-5); 1-IV long, usually 4, 5b (8-5); 1-V very long, usually single

(single, double). Segment VIII: sclerotized plate large but not ringing segment;

anterior comb scales usually 19-21 (18-27), posterior comb scales usually 7-

10 (5-12); larger anterior comb scales shaped like posterior comb scales, with

1 strong but short apical spine and a fringe, the distal members of the fringe

enlarged, about half as long as terminal spine; posterior row of comb scales

about 0.46-0.77 length of anterior row; hair 3-VIII usually 10-12b (9-14). Si-

phon: index usually 4.0-6.0 (3.6-6.17); base of siphon dark brown to black,

- portion between base and alveolus of hair 1-S tan to brown, portion between 1-S

and apex darker, dark tan to dark brown, apex straw-colored to yellowish-tan;

weakly to moderately developed imbricate integumentary sculpturing present in

basal portion; hair 1-S located 0.32-0.40 from base of siphon, usually 9-11b

(6-14). Anal Segment: base and dorsoapical portions usually brown, remainder

usually tan in color; integumentary sculpturing absent or very weakly developed,

imbricate; dorsal anal gills about 1.3-1.9 times length of ventral pair and 1.1-

2.9 times length of anal saddle.

BIONOMICS. The immature stages of albipes are usually found in bamboo

internodes with moderate to large holes or cracks. According to Macdonald

(1958:124, 125), this is the most common species of Orthopodomyia at Ulu Gom-

bak, Malaya, where it is the only one biting man (7 separate records).

SYSTEMATICS. The most distinctive adult feature of this species is the

palpus of the female. It is 5-segmented, with segment 4 moderately long and

segment 5 short. Females of other species in the Albipes group have either a

4-segmented palpus ora 5-segmented one with segment 4 shortand 5 just a minute

apical projection. The added length of segment 4 in albipes effectively separates

the light scales at the base and apex of the segment into 2 patches, so that the

palpus has 3 white patches beyond the base. In other species it has only 2.

Both sexes are distinct in the ornamentation of hindtarsal segment 2, which has

a broader area of white scales at the base than at the apex. Wing vein 1A is

white-scaled at the base in albipes and this serves to separate the species from

all others except for some specimens of the Anopheloides subgroup.

The male genitalia of albipes are similar to wilsoni, but are easily told

from it by a patch of specialized mesal bristles. These specialized bristles

are also found in madrensis, but that species is distinct from albipes in the

number of strong bristles on the basal mesal lobe and the shape of the aedeagus.

Besides the feature given in the key, the pupa is distinctive in hairs 3, 7-C

both being long and multiple. The size and type of branching of hair I-II is al-

so diagnostic. The larval chaetotaxy is quite variable, hairs 1-M, T, III, IV and

13-T, II-V often being developed as in the Anopheloides subgroup.

This species is known to hybridize with anopheloides. The hybrids, 4 males

from northern Thailand, resemble albipes in the adult stage except that they

have a patch of scales on each side of the clypeus. Larvae and pupae are also

like albipes, but show some less conspicuous features of anopheloides. The

male genitalia of these 4 specimens are very distinct, however, the aedeagus

being unlike that of any species in the group; it is somewhat intermediate in

shape between albipes and anopheloides.

Zavortink: Genus Orthopodomyia 137

Although it is probably distantly related to wilsoni, albipes is distinct enough

on larval and pupal features to be placed in a separate subgroup.

Albipes

DISTRIBUTION. Eastern India to South Vietnam, south to the Malay Pen-

insula and northern Borneo. Material Examined: 226 Specimens; 250, 26 9,

28 P, 147 L; 27 individual rearings (22 larval, 4 pupal, 1 incomplete).

FEDERATION OF MALAYSIA. Perak: Batang Padang, 8 Mar 1925, H.M.

Pendlebury, 1 & [BMNH]. |

Sabah: Keningau, 11 Aug 1956 (US 56), 2 2 [BMNH]. Tambunan, 1949, D. H.

Colless, 1 2 [BMNH].

Sarawak: Kuching, 2 o° [BMNH].

Selangor: Kuala Lumpur, 8 Apr 1903, & lectotype [BMNH]; 10 Dec 1903,

1? [BMNH]; 1? [BMNH]. Ulu Gombak, Feb 1956 (04), 31po (04-7, 12, 16),

2 1p? (04-8, 14) [LIVER]; 8 Oct 1957 (0352), 1 1p? (0352-5) [LIVER]; same data

(0353), 21po’ (0353-3, 8), 4 1p? (0353-2, 4, 5,7) [LIVER]; 28 May 1958 (0631),

1 lp (0631-3), 1 1p? (0631-4), 19 [USNM]. Ulu Klang jungle, 21 Mar-18 Sept

1903, 70’ [BMNH]. Ulu deel 6 May 1958 (0600), 1 1pco (0600-2), 11p?

(0600-4), 1 1p (0600-3) ) [USNM]; 10 June 1958 (0646), 1 9 [USNM].

: Ginting Simpab, A.T. Stanton: Io, iP, tL (BMNe),

No locality or date: H.P. Hacker, 4 9 [BMNH].

INDIA. Bihar: Sukna, Aug 1928, Sobha Ram, 1°, 1 2? [BMNH].

THAILAND. Kanchanaburi: Khan Salak Phra, 1965 (00383), 1 1p? (00383-

10) [USNM]; same data (00387), 1 1po (00387-1), 1 po’ (00387-100) [USNM];

same data (00389), 1 lpo (00389-1), 1 lp? (00389-2), 1 p? (00389-101), 6L

[USNM]; same data (00392), 1 po (00392-100), 2 L [USNM].

Ranong: Kra Buri, 1964 (RN 72), 2 1p? (72-40, 42) [USNM].

No locality or date: (00383), 2 L [USNM]; (00123), 4 L [USNM]; (00254),

9L [USNM]; (00256), 4L [USNM]; (00259), 5L [USNM]; (00260?), 1 L [USNM];

(00262), 91. [USNM]; (00267), 221 [USNM]; (00287), 8L [USNM]; (00307), 1L

[USNM]; (00310), 6 L [USNM]; (00313), 2 L [USNM]; (00378), 1 L [USNM]:

(00381), 6 L [USNM]; (00382), 1 L [USNM]; (00384), 2 L [USNM]; (00394), 1 L

[USNM]; (00395), 11 L [USNM]; (00399), 3 L [USNM]; (00400), 1 L [USNM];

(00401), 5 L [USNM]; (T 1794), 2.L [USNM]; (NV 19), 1 L [USNM]; (RN 27),

1 po’ (27-8-113) [USNM]; (RN 69), 8 L [USNM].

Additional records from the literature. SOUTH VIETNAM. Cochin China,

L,A (Borel, 1930a:298-302).

Hybrids

DISTRIBUTION. Northern Thailand. Material Examined: 12 specimens;

4, 4P, 4L; 4 larval individual rearings.

THAILAND. Chiang Rai: Doi Sam Sao, 1965 (00332), 4 1p (00332-1-4)

[USNM ].

Additional records from the literature. None.

ANOPHELOIDES SUBGROUP

PUPAE. Cephalothorax: straw-colored to light tan ventrally, tan to light

brown dorsally; integumentary sculpturing absent; 6-C cephalad of 7-C. Trum-

pet: largely brown to dark brown in color with base and apex lighter; broadening

138 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

gradually from base; pinna large. Metanotum: uniformly tan to light brown or

with haltere case and sometimes mesal portion lighter; integumentary sculptur-

ing absent; fourth pair of hairs never developed. Abdomen: anterior segments

tan to light brown in color, posterior segments yellowish-tan to tan; anterior

portion and posterior edge of anterior segments darker; integumentary sculp-

turing moderately developed on anterior segments; hair 1-II moderately to

strongly developed and with mostly simple branches, some much finer than oth-

ers, arising from the base; hair 2-II fine and mesad of 5-II; 5-II long and sin-

gle; 2-IlI, IV fine; 5-V, VI short, extending nearly to or slightly beyond alveolus

of hair 4 of first following segment in non-hairy forms, but long and extending

nearly to or to alveolus of hair 4 of second following segment in hairy forms;

o-VI usually 3-7b (2-8); 9-VII 7-11b and with longest branch reaching 0.6-0.9

distance to alveolus of 9-VIII. Terminal Segments: male genital lobe projecting

considerably beyond median caudal lobe. Paddle: light straw-colored; longer

than broad; external buttress nearly straight in middle portion and with mar-

ginal spicules; midrib convex mesally.

LARVAE. Head: slightly broader than long; integumentary sculpturing ab-

sent; largely tan to dark brown colored, lighter cephalad and around imaginal

eye, collar very dark brown; mental plate tan to brown, usually with 8-10 teeth

on each side; hairs 4-10-C not all equally developed; 13-C moderately devel-

oped, 4-6b (4-7). Antenna: short, largely brown to dark brown in color, usu-

ally lighter towards apex. Thorax: epidermal pigment strongly developed, red

to purple; spicules absent; hairs 0,1,3-7-P not all equally developed. Abdomen:

segment VII with large sclerotized plate; hairs 6-I, I] with individual branches

finer than 6-III, tapering to fine apex and barbed from near base to near apex.

Segment VIII: anterior comb scales fringed only, or with distal elements of

fringe lengthened and spine-like; larger posterior comb scales with 3-5 large

apical spines, the central one of which may be much longer and stronger than

the others, bordered on each side by numerous lesser spines which become

smaller and fringe-like proximally. Siphon: base of siphon dark brown to black,

portion between base and alveolus of hair 1-S tan to brown, portion between 1-S

and apex almost always darker, dark tan to dark brown, apex straw-colored to

tan; integumentary sculpturing absent to very weakly developed, imbricate. Anal

Segment: saddle largely brown in color, dorsal surface and sometimes base

darker; integumentary sculpturing weakly developed, imbricate.

BIONOMICS. Larvae and pupae have been collected in treeholes, bamboo

stumps and artificial containers. Two species included in this subgroup, anoph-

eloides and andamanensis, are often found together.

SYSTEMATICS. This subgroup contains the 2 widespread species andama-

nensis and anopheloides and the geographically restricted papuensis. It is well

defined on larval and pupal characters, but adults of the included species are

Similar to albipes. and cannot always be separated from siamensis.

| This is the most recent, widespread and dominant subgroup in the Albipes

group. It may well have had a hybrid origin, with putative ancestors in the Al-

bipes and Flavicosta subgroups. Such an interpretation is suggested by the fol-

lowing features of the immature stages: (1) the form of the larval comb scales,

(2) resemblance of the hairy larval and pupal forms of this subgroup to albipes,

and (3) the occurrence of immature stages commonly in both treeholes and bam-

boo. If one of the ancestral species was similar in adult ornamentation to the

extant species albipes or siamensis, then the discordance between classifica-

tions based on adults and immature stages might be partially explained.

Some aspects of the larval morphology, such as the shape of the head and

Zavortink: Genus Orthopodomyia 139

antennae, color of the sclerotized parts, and the modal condition of the chaeto-

taxy, are very Similar to the Signifera group. The European member of that

group, pulchripalpis, has several features in the adult which may have been ob-

tained by introgression with the Albipes group. Possibly, then, some features

of the larva of the Anopheloides subgroup have been obtained from the Signifera

group by introgression.

_ Difficulty may be encountered with the key couplet separating the hairy and

non-hairy forms of larvae and pupae of this subgroup because various interme-

diates occur. Semihairy pupae may often be identified to species by the same

characters as non-hairy ones, but identification of the hairiest individuals is

not possible. i

DISTRIBUTION. Same as for the Albipes group.

22. Orthopodomyia papuensis, n.sp.

Figs. 35, 36

TYPES: Holotype: & with associated larval and pupal skins (384-3), Finsch-

hafen, Territory of New Guinea, larva found in a treehole, 28 Nov 1944, C.S.

Lauby [USNM]. Allotype: 2 with associated larval and pupal skins (384-1), same

data as holotype [USNM]. Paratypes: 1 lpo (384-2), 5L, same data as holotype.

Orthopodomyia andamanensis (in part) of Knight and Mattingly (1950:15-16);

Bonne-Wepster (1954:28-30); Stone, Knight and Starcke (1959:122).

Orthopodomyia anopheloides var. andamanensis of Brug (1934:518-519); Knight,

Bohart and Bohart (1944:14, 66, in part); Lee (1944:36); Brug and Bonne-

Wepster (1947:183, in part). ;

FEMALE. Wing: 3.87 mm. Proboscis: 2.48 mm. Forefemur: 2.60 mm.

Abdomen: about 2.8 mm. Differs from siamensis as follows. Head: torus as

dark as or lighter than head capsule or clypeus; decumbent scales all white or

sometimes yellow laterally; light scales on labium cream-colored or white, in

broad ring near middle and large preapical dorsal patch, or sometimes with

preapical patch continued ventrally to form a ring and extending to middle ring

along ventral surface; palpus 0.37 of proboscis, 5-segmented, segment 4 short,

segment 5 minute, white scales on segment 4 also covering segment 5; torus

scales white or dingy. Thorax: postnotum sometimes lighter laterally; acros-

tichal scales white or brown at caudal end, remainder mostly white but with

some brown and/or yellow; dorsocentral scales mostly white at caudal end,

remainder yellow, white and brown; lateral prescutellar scales sometimes

golden in middle portion of row; posterior fossal scales all white or white and

yellow; fossal scales mixed yellow and brown; supraalar scales all white or

i ee ee ee

white to cream-colored; ssp scales in a moderately long row; stp sometimes

with additional bare area below ssp scales. Legs: forecoxal scales white;

preapical brown and light femoral rings not always distinct, especially on an-

terior surface; posterior surface of forefemur light-scaled at base; midfemur

with posterior surface sometimes largely brown-scaled; ventral surface of fore-

and hindtibiae largely cream-scaled; tarsal segment 1 usually white-scaled at

apex; fore- and midtarsal segments 2 and 3 sometimes light-scaled at baseonly,

segments 4 and 5 dark-scaled, light-scaled, or light-scaled only at ends; hind-

tarsal segment 2 with moderately broad white-scaled apical ring and with poste-

140 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

rior and ventral surfaces largely pale-scaled, segments 3 and 4 all white-scaled.

Wing: costal vein probably as in siamensis, but basal patch absent in the few

specimens available, and prehumeral and humeral light patches either absent

or confluent with presectoral; vein Rj sometimes without subcostal light patch;

vein 1A sometimes with white scales at base.

MALE. Similar to female; differs from siamensis as follows. ProboScis:

preapical light-scaling either a broad dorsal patch or a very broad ring. Pal-

pus: about 0.72 of proboscis. Antenna: integument of torus not always lighter

laterally.

MALE GENITALIA (fig. 35). Segment VIII: tergite lobe about as broad as

long, apex emarginate and with small teeth. Segment IX: tergite with 0-3 bristles

on each side. Sidepiece: basal mesal lobe with 4 or 5 stout bristles and I or 2

finer; each stout bristle only slightly longer than the 1 dorsad of it; mesal sur-

face weakly sclerotized distad of basal mesal lobe; specialized mesal bristles

absent, 1 or 2 specialized tergomesal bristles sometimes present. Clasper:

not strongly curved in distal 0.25. Aedeagus: weakly to moderately sclerotized;

central portion with sides nearly parallel or slightly diverging apically, base _

rounded, apex pointed; 1 pair of sternal teeth present about 0. 65-0. 75 distance

from base; without projection between or below ventral parameres. Proctiger:

paraproct sclerotization with 2-4 apical teeth; cercal setae 2-4.

PUPA (fig.35). Abdomen: 3.22mm. Trumpet: 0.55mm. Paddle: 0.64mm.

Cephalothorax: hair 1-C moderately developed and usually double (2-4b); 2, 3-C

strongly to very strongly developed, usually 3,4b, but up to 14b; 4-C strongly

to very strongly developed, usually 2-4b, but up to 8b; 5-C strongly to very

strongly developed, usually 2,3b (2-4); 7-C very strongly developed, usually

double, but up to 5b; 8,9-C moderately to very strongly developed, usually

2,3b, but up to 5b. Metanotum: hair 10-C moderately developed, usually 6-10b

(6-16); 11-C moderately developed, usually 2-4f; 12-C strongly to very strong-

ly developed, usually 3-5b (3-6). Abdomen: hair 1-II moderately developed,

usually 10-20b (7-25); 1-III usually 10-16b (8-20); 5-IV 4-7b; 5-V, VI 3-6b,

extending at least 1.5 distance to alveolusof hair 4 of second following segment;

9-VIII 9-13b and with longest branch reaching about 0.72 distance to apex of

paddle. Paddle: asymmetrically obovate in outline, external portion sometimes

somewhat angular.

LARVA (fig. 36). Head: 1.20mm. Siphon: 1.31mm. Anal Segment: 0.44mm.

Thorax: 8-P 5-7b; 1-M very long, single; 1-T long, single; 13-T moderately to

strongly developed, 4-7b. Abdomen: hair 1-I, II moderately to very long, usu-

ally single (single, double); 6-I, Il usually 5-8b; 1-III-V long to very long, sin-

gle; 13-III,IV long, single or double, when double 1 branch much longer than

other; 13-V moderately long to long, usually 2,3b. Segment VIII: sclerotized

plate large, sometimes ringing segment; anterior comb scales usually 24-26

(23-27), posterior comb scales usually 9-12 (7-12); posterior row of comb

scales about 0. 64-0. 90 lengthof anterior row; hair 3-VII usually 9-14b. Siphon:

index 5.6-9.5; hair 1-S located 0.33-0.39 from base of siphon, usually 9-13b.

Anal Segment: dorsal anal gills 1.5-1.8 times length of ventral pair and about

1.0-1.1 length of anal saddle.

BIONOMICS. Larvae have been collected in holes in buttress roots, logs

and trees.

SYSTEMATICS. Each stage of papuensis is very similar to at least 1 other

species in the Albipes group. The adults cannot be completely separated from

andamanensis, anopheloides or siamensis and the male genitalia are like sia-

mensis. The pupa is apparently indistinguishable from andamanensis and the

~Zavortink: Genus Orthopodomyia 141

larva from anopheloides. Papuensis is being described as new because the

male genitalia are unlike those of the other 2 species, andamanensis and anoph-

eloides, in the Anopheloides subgroup. Knight and Mattingly (1950:15-16) did

not distinguish papuensis from andamanensis.

The adults have some characteristics which may prove to be diagnostic

when additional specimens are available. Some segments of the legs, particu-

larly the ventral surface of the fore- and midtibiae and the posterior and ven-

tral surfaces of hindtarsal segments 1 and 2, seem to have lighter scales than

andamanensis or anopheloides. The mesonotal scales may prove to be consis-

tently whiter in papuensis than these other species also. |

Most of the pupae available for study are hairy forms. The hairs are usu-

ally more highly branched but shorter than they are in the hairy forms of anda-

manensis and anopheloides. Some of the larvae display a tendency towards

hairiness as well. ©

_ Papuensis, restricted, as far as is definitely known, to Ceram and New

Guinea, is the only allopatric species in the Albipes group. I believe that it is

a relict predating the evolution of andamanensis and anopheloides which has

persisted only on these outlying islands.

DISTRIBUTION. Ceram and New Guinea; possibly also in northern Queens-

land. Material Examined: 41 specimens; 4c, 92, 9 P, 19 L; 9 larval individ-

ual rearings. ,

INDONESIA. Ceram. Karloetoe Kara, 30 Dec 1931, S.L. Brug and H. de

Rook, 1 lpo" (6639), 2 1p? (2898, 6642), 49, 3L [UCLA], 1 lpo (2896), 1 1p?

(2897), 1 L [BMNH]. |

West Irian: Sukarnapura, Pegunungan Cycloop, 21 Feb 1945, Schultz, L. E.

Rozeboom and J. L. Laffoon (270), 1 lp (270.8), 1 L [USNM].

TERRITORY OF NEW GUINEA. Finschhafen, 28 Nov 1944, C.S. Lauby

(384), lpo holotype (384-3), 1 lpo (384-2), 1 1p? (384-1), 5 L [USNM].

Additional records from the literature. AUSTRALIA. Queensland: north-

ern portion, 1L (Marks, 1962, in litt.). The specimen reported as andamanen-

sis may be papuensis.

23. Orthopodomyia andamanensis Barraud

Figs. 37, 38

1934. Orthopodomyia anopheloides var. andamanensis Barraud, 1934:102.

*TYPE: lectotype % with genitalia slide, specimen labeled ''Type °,"

Andaman Islands, Aug 1926, G. Covell; PRESENT SELECTION [BMNH].

Elevated to specific rank by Knight and Mattingly (1950:15-16); reduced

to variety of anopheloides by Thurman (1963:55); resurrected by Delfinado

(1966:68). :

Orthopodomyia andamanensis of Knight and Mattingly (1950:15-16, in part);

Bonne-Wepster (1954:28-30, in part); Macdonald (1957:14; 1958:123, 125, in

part); Stone, Knight and Starcke (1959:122, in part); Macdonald and Traub

(1960:94, in part); Stone (1961:36); Delfinado, Viado and Coronel (1962:438);

Delfinado (1966:68); Stone, Scanlon, Bailey, Delfinado and Bram (1966:82).

Orthopodomyia (Orthopodomyia) andamensis of Thurman (1959:58).

Orthopodomyia anopheloides var. andamanensis of Knight, Bohart and Bohart

(1944:14, 66, in part); Baisas (1946:33, 35, 39,41, 45); Brug and Bonne-Wep-

ster (1947:183, in part); LaCasse and Yamaguti (1948:270); Thurman (1963:

55), ,

2 dag Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

Orthopodomyia albipes of Leicester (1908:176-177, in part).

FEMALE. Wing: 5.35 mm. Proboscis: 2.86 mm. Forefemur: 3.13 mm.

Abdomen: about 4.2 mm. Differs from papuensis as follows. Head: decumbent

scales all white; posterior dorsolateral, lateral and ventral scales white; labi-

um with light scales usually white, with ring either at or distad of middle or ab-

sent, with dorsal preapical patch sometimes either absent or continued ventral-

ly to form a ring, and sometimes with additional light scales scattered near

base; palpus 0.43 of proboscis, sometimes with white scales at joint between

segments 1-2; torus scales white. Thorax: acrostichal scales usually brown,

sometimes yellow, at caudal end, remainder sometimes largely yellow and ©

white but with some brown; dorsocentral scales white or white and yellow at

caudal end, remainder yellow and/or cream-colored and brown; lateral pre-

scutellar scales rarely golden laterally; supraalar scales entirely white or

white mesally, and sometimes at ends of anterior and middle projections, the

remainder yellowish, yellow or golden; ppn with all scales narrow and curved

or lower ones broad and flat, lowermost and usually uppermost scales white,

remainder yellow. Legs: forecoxal scales white to cream-colored; preapical

femoral rings usually distinct; forefemur sometimes with a small light-scaled

patch basad of preapical brown-scaled ring on anterior surface, and with basal

scaling of posterior surface dark or light; midfemur not always brown-scaled

to beyond middle on anterior surface and posterior surface with light-scaling

yellowish or cream-colored; mid- and hindtibiae sometimes with a more or

less conspicuous largely white- to yellowish-scaled ring distad of subbasal —

brown-scaled ring and foretibia usually with an anterior light patch in same

position, anterior surface of mid- and hindtibiae with either light or dark scales

predominating; ventral surface of fore- and hindtibiae brown-scaled with light

speckles; foretarsus entirely dark-scaled or segments 2 and 3 white-scaled at

base and sometimes at apex and segments 4 and 5 with a few white scales at

base; midtarsal segment 2 and usually 3 white-scaled at both ends, segments 4

and 5 white- to light brown-scaled; hindtarsal segment 2 with narrow to mod-

erately broad apical white ring and with posterior and ventral surfaces largely

dark-scaled, segments 3 and 4 entirely white-scaled or rarely with moderately

to very broad subbasal dark-scaled ring on segment 3. Wing: costal vein usu-

ally with accessory subcostal light patch and usually with distinct basal, prehu-

meral, humeral and presectoral patches, or 1 or more of basal, prehumeral,

humeral and presectoral patches absent, or humeral and presectoral patches

large and confluent; vein Rj rarely lacking accessory subcostal patch. Haltere:

scales usually transparent to whitish, sometimes some brownish. Abdomen:

tergite I fan sometimes with white scales; light scales usually white or cream-

colored, rarely yellowish; tergites II-V sometimes with light scales basally.

MALE. Similar to female; differs from papuensis as follows. Proboscis:

always with white or rarely yellowish scales in a moderately broad to broad

preapical ring which extends further basally along ventral surface. Palpus:

about 0.80 of proboscis. Abdomen: tergites and sternites sometimes with light-

scaled basal bands; sternite VIII light-scaled basally, laterally and apically.

MALE GENITALIA (fig. 37). Segment VIII: tergite lobe slightly to consid-

erably longer than broad, apex usually emarginate, sometimes truncate, and

with small teeth. Segment IX: tergite with 0-2 bristles on each side. Sidepiece:

basal mesal lobe with 4 or 5 stout bristles and 1-3 finer; each stout bristle only

slightly longer than the 1 dorsad of it; mesal surface usually weakly sclerotized

distad of basal mesal lobe; specialized mesal bristles absent, 1 or 2 specialized

Zavortink: Genus Orthopodomyia 143

tergomesal bristles present. Clasper: not strongly curved in distal 0.25. Ae-

deagus: weakly sclerotized; broadest distally, tapering to obtuse apex, sides

almost parallel basad of broadest point but with constriction near middle; with-

out sternal teeth but with basal projection that extends between and below ven-

tral parameres, apical portion of projection usually with pair of small teeth;

major direction of basal projection is below ventral parameres. Proctiger:

paraproct sclerotization with 3 or 4 apical teeth; cercal setae 3 or 4.

PUPA (fig.37). Abdomen: 3.72 mm. Trumpet: 0.56 mm. Paddle: 0.86

mm. Cephalothorax: hair 1-C moderately developed and usually double (2, 3b)

in non-hairy form, strongly developed and up to 4,5b in hairy form; 2,3-C

moderately developed and usually 3-5b (2-6) in non-hairy form, strongly to very

strongly developed and up to 9,10b in hairy form; 4-C moderately to strongly

developed and 2-4b in non-hairy form, very strongly developed and up to 7, 8b

in hairy form; 5-C strongly to very strongly developed and 2, 3b in non-hairy

form, very strongly developed and up to 6b in hairy form; 7-C very strongly

developed, 2,3b in non-hairy form, 5, 6b in hairy form; 8, 9-C moderately to

strongly developed and usually 2, 3b (2-5) in non-hairy form, very strongly de-

veloped and up to 7, 8b in hairy form. Metanotum: hair 10-C moderately devel-

oped, usually 10-13b (10-17); 11-C moderately developed, usually 2-4f; 12-C

strongly developed, usually 5, 6b (5-9). Abdomen: hair 1-II moderately devel-

oped and with 16-25 branches in non-hairy form, strongly developed and with

up to 35 branches in hairy form; 1-III usually 12-14b (10-17); 5-IV 5, 6b in non-

hairy form, up to 8b in hairy form; 5-V usually 4, 5b (4-6) in non-hairy form,

up to 6, 7b in hairy form; 5-VI usually 6, 7b (4-8); 9-VIII 12-16b and with long-

est branch reaching 0.6-0.7 distance to apex of paddle. Paddle: asymmetrical-

ly and narrowly to broadly obovate in outline, external portion somewhat angu-

lar.

LARVA (fig.38). Head: 1.24 mm. Siphon: 1.57 mm. Anal Segment: 0.52

mm. Thorax: 8-P usually 9-11b (8-12); 1-M very long, single; 1-T long, sin-

gle; 13-T moderately to strongly developed, usually 5-9b. Abdomen: hairs 1-I,

II moderately long, usually single (single, double); 6-I usually 9, 10b (7-10), 6-

II usually 9-11b (7-12); 1-III, V very long, single; 13-III, IV long, usually dou-

ble (1-3b), one branch much longer than other; 1-IV long, usually single (sin-

gle, double); 13-V moderately long, usually 3, 4b (2-5). Segment VIII: sclero-

tized plate large, sometimes ringing segment; anterior comb scales usually

24-28 (22-30), posterior comb scales usually 10-12 (8-14); posterior row of

comb scales about 0. 69-0. 85 length of anterior row; hair 3-VIII usually 10-14b.

Siphon: index usually 4.0-5.6; hair 1-S located 0.37-0.50 from base of siphon,

usually 11,12b (10-14). Anal Segment: dorsal anal gills 1.5-2.0 times length

of ventral pair and 1.2-2.2 length of anal saddle.

BIONOMICS. Knight and Mattingly (1950:16) report the capture of a female

from Celebes biting in the daytime. Nothing else is known of the habits of the

adults. The entire range of breeding sites of the Anopheloides subgroup is uti-

lized by andamanensis.

SYSTEMATICS. The decision to consider andamanensis as a species dis-

tinct from anopheloides is a very tenuous one. The 2 forms are extremely sim-

ilar morphologically in all stages, appear to have the same ecology and are

sympatric over a broad geographical area. It is possible that andamanensis is

only a form of anopheloides in which certain slight adult, larval and pupal vari-

ations are correlated. Analysis of the 2 forms has been hampered by the pau-

city of large series of individuals and associated larval and pupal skins. A final

decision on the status of andamanensis cannot be reached without further collec-

144 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

tions, ecological studies and progeny rearings. Although doing socreates prob-

lems, I favor the hypothesis that the 2 forms are distinct species and am treat-

ing them as such here.

Andamanensis and anopheloides can be separated more reliably in the adult

stage than in any other. Andamanensis has a very narrow to moderately broad

white ring at the apex of hindtarsal segment 2 and usually has hindtarsal seg-

ments 3 and 4 entirely white-scaled, whereas anopheloides has a narrow to very

broad white ring at the apex of hindtarsal segment 2 and usually has a dark ring

or patch distad of the middle of tarsal segment 3 or segments 3 and 4. The api-

cal white ring of hindtarsal segment 2 is usually less than 2 times, but up to 3

times, as long as the diameter of that tarsal segment in andamanensis; in

anopheloides the ring is never narrower than 2 diameters and is usually greater

than 3. There are 3 male specimens from Thailand which have dark scales in

a ring in the basal portion of hindtarsal segment 3. I am referring these to an-

damanensis on the basis of the male genitalia and the very narrow white ring on

the apex of hindtarsal segment 2. These 3 adults can be separated from those

of anopheloides which have the dark scales restricted to hindtarsal segment 3

because the dark-scaling is distad of the middle of the segment in that species.

They could possibly represent a new species, but in the absence of immature

stages it seems better to consider them as andamanensis. Rare individuals of

anopheloides have hindtarsal segments 3 and 4 entirely white-scaled. These

specimens are the most difficult to separate from andamanensis, but can some-

times be distinguished by a broader white ring at the apex of hindtarsal segment

2. Judging by the broad apical ring on hindtarsal segment 2, the leg of ''anda-

manensis" illustrated by Macdonald (1958:122) is most likely that of a specimen

of anopheloides with the last 3 hindtarsal segments entirely white.

Separation of the male genitalia of andamanensis and anopheloides is diffi-

cult. The only way to differentiate between them is by the direction of the pro-

jection from the base of the aedeagus. In andamanensis the projection is usual-

ly directed more sternad than cephalad (towards segment VIII), while the re-

verse is usually true in anopheloides. However, there are specimens which

appear to be one species on the basis of leg banding which have the aedeagus of

the other species. In addition, the angle of the aedeagus, and hence of the pro-

jection, varies from specimen to specimen due to the varying degree to which

the aedeagus may be swung forward by the ventral parameres, and, as a con-

sequence, the apparent major direction of the projection may not be its direc-

tion in the resting position.

Separation of the immature stages of the 2 species is also difficult. In the

collection of W. W. Macdonald there are several individual rearings of anda-

manensis and many of anopheloides. The larvae and pupae of these specimens,

all collected near Kuala Lumpur, Malaya, can be separated without much diffi-

culty. Larvae of andamanensis differ from anopheloides by a longer and darker

siphon, hair 1-S nearer the base of the siphon, fewer branches in hair 0-P, a

greater number of branches in hairs 6-I, II and a greater number of posterior

comb scales. Pupae of andamanensis differ by a broader outer part of the pad-

dle and a greater number of branches in hairs 9,10-C and 1,6-II]. Examination

of associated skins from other localities where the 2 species have been collec-

ted in the same treehole or in the same immediate area shows that only a part

of these same differences are present. On the average, the andamanensis lar-

‘va seems to have always a longer siphon and a greater number of branches in

hairs 6-I, II and the pupa seems to have always a greater number of branches

in hairs 9,10-C and 1-III. However, the length of the siphon (and the siphon

Zavortink: Genus Orthopodomyia 145

index) and the number of branches in the hairs involved for both the larva and

pupa vary to such a degree that it is not possible to achieve reliable separation

of the species by a key. Specimens of the immature stages from an area from

which no associated skins are available for comparison often cannot be identi-

fied. The most dependable hair branchings for separation of the associated

skins available are given in the Key.

One collection from Kuala Lumpur has non-hairy larval and hairy pupal

skins associated with both anopheloides and andamanensis adults. This can be

interpreted in 1 or 2 ways. It is either good evidence that hairy forms are en-

vironmentally induced variations or that andamanensis is indeed only a form of

anopheloides. On the basis of observations made on other species (see arbor-

icollis) the first alternate is favored. Hairy larval forms have not been seen

for andamanensis, but they are to be expected.

DISTRIBUTION. Eastern India to South Vietnam and Palawan, south to

the Andaman Islands, Sumatra, Java and Celebes. Material Examined: 145

specimens; 560, 4492, 1 A, 1 isolated % genitalia, 20 P, 23 L; 16 individual

rearings (12 larval, 2 pupal, 2 incomplete).

FEDERATION OF MALAYSIA. Perak: Pulau Pangkor Laut, 19-23 Oct

1903, C.W. Daniels, 2 2 [BMNH].

Selangor: Ulu Gombak, 25 Oct 1956, J.A. Reid (084), 1 1po (084-29) [LIVER];

13 Nov 1956 (096), 1 1p (096- -38) ) [LIVER]; 10 Jan 1957, J.A. Reid (0129), 1 lpo&

(0129-27), 3 1p? (0129-22, 30, 41) [LIVER], 1 1p (0129- 24) [USNM]; 15 Jan 1957,

J.A. Reid (0133), 10, 12 [USNM]; 14 Feb 1957 (0175), 1% [LIVER].

INDIA. Andaman and Nicobar Islands: Andaman Islands, Aug 1926, G.

Covell, o lectotype, 1 2 [BMNH].

INDONESIA. Java: Bogor, 1930, R.W. Paine, 1P [BMNH]. Garut, Djaja-

sana, 1 9 [UCLA].

Kalimantan: Samarinda, 1 1po’ (9711), 19 [UCLA]. Tarakan, 12 June 1945,

A.G. Humes (30), 1 o [USNM].

Sumatra: Danau Ranau, 50°, 62, 1P, 1L [UCLA]. Tjoeroek, 1? [UCLA].

REPUBLIC OF THE PHILIPPINES. Palawan: Irahuan River area, 2 June

1945, J. L. Laffoon (854), 1 lpo (854.25), 1 lp? (854.30) [USNM].

SINGAPORE. D.H.C. Given, 2°, 19, 1 P, 1 L [BMNH].

SOUTH VIETNAM. Long Khanh, 1964 (LK 2746), 1 2 [USNM]. Long Khanh,

Xuan Loc, 1965 (LK 2781), 1 o [USNM].

THAILAND. Chiang Mai: Chang Puek, 1962 (T 1041), 10 [USNM]. Doi

Suthep, 1962 (T 215 E), 12 [USNM]; same data (T 1095-1), 1 2 [USNM]; same

data (T 1147), 1 o& [USNM]; same data (T 1161), 1 [USNM]; same data (T

1185), 3 o [USNM]; same data (T 1224), 2 o [USNM]; same data (T 1384), 1 o

[USNM]; same data (T 1659), 2, 1? [USNM]; same data (T 1926), 10, 19

[USNM]; 1963 (T 1934), 3 o [USNM]; same data (T 1936), 10, 1 9 [USNM];

same data (T 1989), 1 o% [USNM]. Huay Mae Sanan, 1962 (T 1157), 50°, 49

[USNM ].

Chiang Kai: Doi Sam Sao, 1965 (00277), 2 po (00277-100, 101) [USNM].

Khon Kaen: Pa Dong Larn, 1962 (T 1861), 1 9 [USNM].

Lampang: Kuhn Tal, 10 June 1952, D.C. i hebman (M 208), 1% genitalia

USNM].

Nakhon Nayok: Khao Kheo, 17 Sept 1963 (NY 43), 1 1p? (43-20) [USNM].

Khao Yai, 23 May 1964 (NY 151), 4L [USNM]. Sariga Village, 22 May 1964

(NY 140), 1 lp? (140-41), 119 (140-43) [USNM].

Nakhon Ratchasima: Khao Yai, 1962 (T 1863), 1 o [USNM].

No locality or date: (00303), P [USNM]; oe 169), 1L [USNM]; (CM 381),

146 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

1 lpo (381-40) [USNM]; (RN 29), 1 P [USNM]; (RN 36), 2c [USNM]; (TG 55),

1 L [JUSNM]; (TG 59), 1 L [USNM]; (T 1225), 3 0°, 22 [USNM]; (T 1226), 1 o

[USNM]; (T 1862), 1 o [USNM]; (T 2094), 1 o [USNM]. |

NO DATA. 60, 99, 1 A [UCLA]. |

Additional records from the literature. INDIA. Bihar: Sukna, Aug 1928,

Sobha Ram, A (Barraud, 1934:102).

Uttar Pradesh: Dehra Dun, L (Wattal, Bhatia and Kalra, 1958:226-228).

I strongly suspect that these larvae are anopheloides rather than andamanensis.

INDONESIA. Celebes: Mamudju, A (Knight and Mattingly, 1950:15-16).

24. Orthopodomyia anopheloides (Giles)

Figs. 39, 40, 41

1903. Mansonia anopheloides Giles, 1903:315. *TYPE: lectotype & with geni-

talia slide, specimen labeled ''Mansonia anopheloides o type,'’ Himalayan

foothills, Dehra Dun, [Uttar Pradesh], India, reared from a larva collec-

ted in a garden tank; PRESENT SELECTION [BMNH].

1908. Orthopodomyia albipes var. nigritarsis Leicester, 1908:177. *TYPE:

lectotype ¢ (76), specimen labeled ''cotype,'' ''The Dinding,"' on the island

Pangkor Laut, [Perak], Malaya, reared from a larva found in a treehole,

20 Oct 1903, Daniels; PRESENT SELECTION [BMNH]. Synonymy with

anopheloides by Edwards (1913:239); synonymy with albipes by Edwards

(1932:108); synonymy with anopheloides by Barraud (1934:98).

1910. Orthopodomyia maculata Theobald, 1910a:29-30. TYPE: holotype ,

Maddathorai, west base of Western Ghats, Travancore, India, adult found

resting on a tree trunk in the jungle, 17 Nov 1908, Annandale [IM]. Syn-

onymy with anopheloides by Edwards (1913:239); considered as a form of

anopheloides by Barraud (1927:527, 529); considered as a synonym of

anopheloides by Edwards (1932:108); considered as a variety of anophe-

loides by Barraud (1934:101); given subspecific. rank by Knight and Mat-

tingly (1950:9-10); elevated to specific rank by Thurman (1959:59); re-

duced to variety of anopheloides by Thurman (1963:55).

1910. Orthopodomyia maculipes Theobald, 1910b:470-473. *TYPE: lectotype

?, specimen labeled as "type 2"' by Theobald, Andaman Islands, 22 July

1908, Lowis and R. White; PRESENT SELECTION [BMNH]. Reduction

to varietal status of anopheloides suggested by Edwards (1913:239); con-

sidered as a form of anopheloides by Barraud (1927:527, 528, 529); resur-

rected by Edwards (1928:53); reduced to a variety of anopheloides by

Barraud (1934:101); resurrected by Knight and Mattingly (1950:10-13).

NEW SYNONYMY.

1946. Orthopodomyia manganus Baisas, 1946:35-39. TYPE: holotype % with

associated larval and pupal skins (TH 111-5), Llavac, Laguna, Luzon,

Philippine Islands, larva found in a treehole, 14 Sept 1940, P. Sunico

[Location unknown]. Synonymy with anopheloides by Knight and Matting-

ly (1950:7).

1948. Orthopodomyia (Orthopodomyia) nipponica LaCasse and Yamaguti, 1948:

264-269. *TYPE: lectotype 2? with associated pupal skin, heavily wooded

cemetery in the foothills of the mountainous terrain surrounding the city

of Kyoto, Honshu, Japan, reared from a larva found in a cut bamboo

flower holder or a stone bowl, 17 Nov 1948; selection of Stone and Knight

(1957:198-199) [USNM]. Synonymy with anopheloides by Knight and Mat-

tingly (1950:7).

Zavortink: Genus Orthopodomyia _ 147

1959. Orthopodomyia (Orthopodomyia) lemmonae Thurman, 1959:58-59. *TYPE:

holotype larva (M398-1), Doi Chom Cheng of Doi Sutep Range, Chiang Mai

Province, Thailand, larva hatched from debris collected in a treehole,

30 Dec 1952, D.C. Thurman [USNM, 64290]. NEW SYNONYMY.

Orthopodomyia anopheloides of Edwards (1913:239); Senior-White (1920:317);

Edwards (1922:458, 470); Barraud (1927:527-529, in part); Barraud and Co-

vell (1928:676); Edwards (1928:53); Barraud (1932:1014-1015); Edwards

(1932:108); Barraud (1934:98-102); Brug (1934:519); Wu (1936:46, 51); Feng

(1938:287; 1939:1581); Edwards (1941:72, 73); Baisas (1946:33, 35, 36, 39);

Bohart (1946:5); LaCasse and Yamaguti (1948:269, 270); Chow (1949:128;

1950:282); Knight and Mattingly (1950:7-9); LaCasse and Yamaguti (1950: ,

65); Kimizu (1952:25-28); Asanuma and Nakagawa (1954:373); Chen and Lien

(1956:204); Chang (1957a:214, 220, 221, 222: 1957b:503-505, 508-509, 510);

Gentry (1957:82); Hara (1957b:16); Macdonald (1957:14); Ramachandra Rao

and Rajagopalan (1957:13, 33); Macdonald (1958:125); Bohart (1959:196);

Stone, Knight and Starcke (1959:122); Macdonald and Traub (1960:94); Del-

finado, Viado and Coronel (1962:438); Lien (1962:621-622); Thurman (1963:

55); Kalra and Wattal (1965:316); Delfinado (1966:66-67); Stone, Scanlon,

Bailey, Delfinado and Bram (1966:82).

Orthopodomyia (Orthopodomyia) anopheloides of Thurman (1959:56-57).

Orthopodomyia (anopheloides) anopheloides of Hara (1957a:50).

Mansonia anopheloides of Giles (1903b:329).

_Finlaya anopheloides of Giles (1904:365, 366); Blanchard (1905:631); Theobald

(1905:33). :

_Orthopodomyia maculipes of Edwards (1913:239; 1922:459, 470); Haga (1924:

831); Edwards (1926:117-118; 1928:53, 57); Brug and Edwards (1931:259);

Edwards (1932:108); Brug and Bonne-Wepster (1947:183); Knight and Mat-

tingly (1950:10-13); Macdonald (1957:15; 1958:123, 125, 126); Stone, Knight

and Starcke (1959:123); Macdonald and Traub (1960:94); Delfinado, Viado

and Coronel (1962:438); Thurman (1963:54); Delfinado (1966:70).

Orthopodomyia (Orthopodomyia) maculipes of Thurman (1959:59-60).

Orthopodomyia anopheloides var. maculipes of Barraud (1934:101); Baisas

(1946:33, 35, 39, 41).

Orthopodomyia anopheloides form maculipes of Barraud (1927:527, 528, 529).

Orthopodomyia maculata of Theobald (1910b:473-474).

Orthopodomyia (Orthopodomyia) maculata of Thurman (1959:59).

Orthopodomyia anopheloides maculata of Knight and Mattingly (1950:9-10);

Stone, Knight and Starcke (1959:122).

Orthopodomyia anopheloides var. maculata of Barraud (1934:101); Baisas (1946:

33, 35, 36); Chu (1957:149, 163; 1958:110,111); Thurman (1963:55).

Orthopodomyia anopheloides form maculata of Barraud (1927:527, 529).

Orthopodomyia lemmonae of Stone (1961:36).

Orthopodomyia (O.) nipponica of Stone and Knight (1957:198-199).

Orthopodomyia anopheloides nipponica of LaCasse and Yamaguti (1950:58-65,

app. I:2); Kimiza (1952:25-26).

Orthopodomyia andamanensis of Macdonald (1958:123, 125, in part); Wattal,

Bhatia and Kalra (1958:226-228); Macdonald and Traub (1960:94, in part).

Orthopodomyia albipes of Bohart (1950:403, 404).

Orthopodomyia mcgregori of Knight and Chamberlain (1948:10); Knight and

Mattingly (1950:13-15); Stone, Knight and Starcke (1959:123); Delfinado,

Viado and Coronel (1962:438); Delfinado (1966:68).

148 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

FEMALE (fig.39). Wing: 3.83 mm. Proboscis: 2.29mm. Forefemur:

2.51 mm. Abdomen: about 3.4 mm. Differs from andamanensis as follows. |

Head: decumbent scales sometimes yellow; palpus 0.48 of proboscis. Thorax:

lateral prescutellar scales brown and white only, sometimes with considerably

more white scales than in andamanensis; ppn sometimes with all scales white;

rarely with remaining pleural scales yellow-tinged. Legs: midfemoral poste-

rior surface sometimes largely dark-scaled; hindtarsal segment 1 sometimes

with broad apical white ring; hindtarsal segment 2 dark-scaled at base or with

a few white scales and with a moderately to very broad white apical ring, seg-

ments 3 and 4 entirely white-scaled, or more commonly segment 3 with brown

scales in small dorsal patch to broad ring distad of middle, or even more com-

monly segment 3 with broad brown ring distad of middle and segment 4 with

brown scales in small dorsal patch to moderately broad ring distad of middle.

Wing: costal vein sometimes with basal, prehumeral and humeral patches large

and confluent; sectoral patch of costal vein sometimes absent; vein R rarely

with sectoral and accessory sectoral patches large and confluent; vein Rj rare-

ly with subcostal patch absent. Abdomen: tergite I fan infrequently with white

scales; subdorsal median patches usually developed only on distal segments;

tergites and sternites sometimes with white basal bands.

MALE (fig.39). Similar to female; differs from andamanensis as follows.

Proboscis: preapical ring sometimes reduced to dorsal patch. Palpus: about

0.78 of proboscis. Antenna: fifth flagellar segment rarely with tuft of scales.

Abdomen: tergites and sternites more frequently and extensively banded than in

female; sternite VIII light basally and laterally.

MALE GENITALIA (fig. 40). Segment VIII: tergite lobe longer than broad,

sides parallel or diverging apically, apex emarginate or truncate and usually

with small teeth. Segment IX: tergite with 1-4 bristles on each side. Side-

piece: basal mesal lobe with 4-6 stout bristles and 1-3 finer; each stout bris-

tle only slightly longer than the 1 dorsad of it; mesal surface membranous or

weakly sclerotized distad of basal mesal lobe; mesal surface without any spe-

cialized bristles or with weakly developed curved or Sinuous specialized bris-

tles distad of basal mesal lobe; 1, 2 specialized tergomesal bristles present.

Clasper: not strongly curved in distal 0.25. Aedeagus: weakly to moderately

sclerotized; as in andamanensis except that major direction of basal projection

is between rather than below ventral parameres. Proctiger: paraproct sclero-

tization with 2-4 apical teeth; cercal setae 3-6.

PUPA (fig.40). Abdomen: 3.44 mm. Trumpet: 0.55 mm. Paddle: 0.92

mm. Cephalothorax: hair 1-C moderately developed and 2,3b in non-hairy

form, moderately to strongly developed and up to 3, 4b in hairy form; 2-C mod-

erately developed and 1-3b in non-hairy form, very strongly developed and up

to 6, 7b in hairy form; 3-C moderately developed and 3-5b in non-hairy form,

very strongly developed and up to 9,10b in hairy form; 4-C moderately to very

strongly developed and 2, 3b in non-hairy form, very strongly developed and up

to 5, 6b in hairy form; 5-C very strongly developed, 2, 3b in non-hairy form, up

to 5, 6b in hairy form; 7-C very strongly developed, double in non-hairy form,

up to 5, 6b in hairy form; 8,9-C moderately to strongly developed and usually

single (sometimes double) in non-hairy form, strongly to very strongly devel-

oped and 1-3b in hairy form. Metanotum: hair 10-C moderately developed,

usually 3-6b (2-8); 11-C moderately developed, usually 3, 4f (2-5); 12-C usu-

ally 2,3b (2-4) and moderately or, especially in hairy form, strongly developed.

Abdomen: hair 1-II moderately developed and usually with 12-19 (8-23) branches

in non-hairy form, strongly developed and up to 34-branched in hairy form;

Zavortink: Genus Orthopodomyia 149

1-TII usually 5-7b (3-11); 5-IV 3, 4b in non-hairy form, up to 7, 8b in hairy

form; 5-V usually 3, 4b (2-4) in non-hairy form, up to 6, 7b in hairy form; 5-VI

usually 3-5b (2-6); 9-VIII 9-14b and with longest branch reaching 0.4-0.9 dis-

tance to apex of paddle. Paddle: usually narrowly and asymmetrically ellipti-

cal or obovate in outline.

LARVA (fig.41). Head: 1.05 mm. Siphon: 1.15 mm. Anal Segment: 0.50

mm. Thorax: 8-P usually 8-12b (5-14); 1-M very long and single in non-hairy

form, long and up to 4b in hairy form; 1-T long and single in non-hairy form,

long and up to 4b in hairy form; 13-T moderately to strongly developed, or, es-

pecially in hairy form, very strongly developed, usually 4-7b. Abdomen: hairs

1-I, II moderately long and single or double in non-hairy form, long and up to

ob in hairy form; 6-I usually 5-8b, 6-II usually 4-8b; 1-III, V very long and sin-

gle in non-hairy form, long and double in hairy form; 13-III, IV moderately long

and usually double (1-3b), with 1 branch longer than other in non-hairy form,

long and up to 5b in hairy form; 1-IV long and usually double (1-3b) in non-

hairy form, long and up to 5b in hairy form; 13-V moderately long and usually

3b (2-4) in non-hairy form, long and up to 5b in hairy form. Segment VII:

sclerotized plate large but not ringing segment; anterior comb scales usually

21-28 (17-32), posterior comb scales usually 7-10 (6-12); posterior row of

comb scales about 0.55-0.74 length of anterior row; hair 3-VIII usually 9-13b.

Siphon: index usually 3.5-4.8 (3.2-6.4); hair 1-S located 0.36-0.50 from base

of siphon, usually 10-12b (9-13). Anal Segment: dorsal anal gills 1.6-2.5 times

length of ventral gills and 0.71-2.5 length of anal saddle.

BIONOMICS. The immature stages may be found in treeholes, bamboo

stumps and artificial containers in association with those of flavicosta, flavi-

thorax and madrensis in addition to those of andamanensis. Adults have been

collected resting on tree trunks.

SYSTEMATICS. This is the most widespread species in the Albipes group,

occupying the entire range of the group except: for the islands east. of Borneo

and south of the Philippines. Because of its broad distribution and variability,

particularly in ornamentation of the proboscis and hindtarsus, the species has

numerous Synonyms.

In addition to anopheloides, 4 names (nigritarsis, maculata, manganus and

nipponica) have been given to adults with dark rings on hindtarsal segments 3

and 4. Leicester apparently did not know of Giles' 1903 description of anophe-

loides when he described nigritarsis in 1908. Although Leicester was the au-

thor of the species albipes, the "'albipes" he described in the 1908 "'Culicidae

of Malaya" was actually andamanensis. The type of nigritarsis differs in no

way from anopheloides and this nominal species has been considered as a syn-

onym of the latter by virtually all authors.

According to Giles (1904:365), Theobald had placed anopheloides with an

allied species (albipes) in the genus Orthopodomyia in the collection at the Brit-

ish Museum. Theobald apparently overlooked or forgot the name anopheloides

later, for it is not mentioned in the 1910 description of maculata or the 1910

volume of A Monograph of the Culicidae. It cannot be determined, then,

whether he described maculata because he was ignorant of anopheloides or be-

cause he felt that it was distinct from anopheloides. At any rate, the names

are applied to individuals which agree in all details except that maculata has a

broader white ring at the apex of hindtarsal segment 2. It has been considered

to be anything from a synonym, form, variety or subspecies of anopheloides to

being a distinct species. Knight and Mattingly (1950:9-10) considered it to bea

subspecies of anopheloides. I am treating it as a synonym of anopheloides,

150 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

considering the differences in hindtarsal banding to be within the limit of var-

iation of that species. Specimens of anopheloides with a broad white ring at the

apex of hindtarsal segment 2 have been seen in collections from Borneo, Cey-

lon, Java, the Philippines, Taiwan and Thailand.

Manganus and nipponica were considered to be distinct from anopheloides

by their authors largely on the basis of ornamentation of the proboscis, but

Knight and Mattingly (1950:9) found both to fall within the range of variation of

anopheloides and reduced them to synonymy. I agree with their interpretation.

The name maculipes was given to a form identical to anopheloides except —

that hindtarsal segment 4 was entirely white-scaled. This nominal species has

been considered to be a form or variety of anopheloides, or a distinct species.

Knight and Mattingly (1950:10-13) treated it as a distinct species. I am reduc-

ing maculipes to synonymy with anopheloides on the basis of specimens from

Malaya in the collection of W.W. Macdonald. As pointed out by Macdonald

(1958:125, 126), the dark band on hindtarsal segment 4 of 1 of his specimens of

anopheloides was incomplete and represented by only a few dark scales. How-

ever, of even more value than this specimen with intermediate tarsal banding

is the large series of adults with associated skins. Comparison of the larval

and pupal chaetotaxy has failed to show any differences between the Malayan

maculipes and anopheloides and has failed to show any differences between the

Malayan anopheloides and anopheloides from near the type locality in India. As

a consequence, it appears that maculipes is just a natural variant of anopheloi-

des and should be considered as a synonym of it. Specimens of anopheloides

with hindtarsal segment 4 entirely white-scaled have been seen in collections

from Malaya, Singapore and Sumatra as well as from the Andaman Islands.

Thurman described 2 fourth instar larvae as lemmonae in 1959, consider-

ing them to be distinct from all other Orthopodomyia species in Thailand be-

cause they lacked the sclerotized basal ring on segment X and the sclerotized

plate on segment VII, had only a small sclerotized plate on segment VIII, and

had the longest branch in hair 2-X bifid, twice the length of other branches and

only 0.33 the length of hair 3-X, whereas all the other species had a sclero-

tized ring on segment X, had large sclerotized plates on segments VII and VHI

and had the longest branch of hair 2-X single, 3 times the length of the shorter

branches and 0.75 the length of hair 3-X. Tate (1932:115) and Reeves (1941:

70-71) had reported that the plates on the terminal segments of Orthopodomyia

larvae were added without moulting by the fourth instar larva and that the num-

ber and size were dependent upon the age of the larva. Since both of Thurman's

larvae died early in the fourth instar, they lack the full complement of plates

simply because they were young. No significance can be attached to hair Ze Xe;

for it may have 1 or 2 branches elongated and the length of these elongated

branches is variable. It is common for the elongated branch to be shorter when

double or bifid than when single. Despite the fact that lemmonae is not differ-

ent on the basis of the characters pointed out by Thurman, its chaetotaxy is

quite different from any species in the Albipes group. Hairs 1-M, T,I-VII and

13-T, INI-V are well developed and multiple. In this respect the larvae resem-

ble albipes, but the morphology of the head, antenna, comb scales and siphon

agree with anopheloides. Another of Thurman's collections contains larvae

identical to the 2 specimens she called lemmonae except that they have only 1

long branch in hair 2-K and segments VII and VIII have sclerotized plates;

these larvae were identified as anopheloides. This was correct, for these lar-

vae, and the 2 she called lemmonae, are only hairy forms of anopheloides.

Similar hairy forms have been seen in collections from Borneo, Japan, the

Zavortink: Genus Orthopodomyia 151

Philippines and Taiwan in addition to Thurman's and others from Thailand. Nu-

merous specimens in collections from Borneo and Thailand show various de-

grees of intermediacy between the normal and hairy extremes and on this basis

lemmonae is reduced to synonymy with anopheloides.

The majority of adults of anopheloides have hindtarsal segments 3 and 4

with a dark ring and relatively few have a dark ring on segment 3 only. In ad-

dition, rare specimens have the last 3 hindtarsal segments all white-scaled. A

specimen from Palawan with this pattern was called mcgregori by Knight and

Chamberlain (1948:10) and Knight and Mattingly (1950:13-15), but is definitely

anopheloides (see mcgregori). Others have been seen from Malaya and possi-

bly the Philippines (Negros) and Sumatra. Means by which these and other

specimens of anopheloides can be told from andamanensis are discussed under |

that species.

Larvae of anopheloides from Japan have longer siphons than those from

other areas. Larvae from the Philippines often have hair 1-T double in other-

wise non-hairy individuals. In addition to hairy larvae, hairy pupae are known.

Normal larvae may give rise to normal or hairy pupae; it is not known if hairy |

larvae give rise to hairy pupae, but that is suspected since partially hairy lar-

vae from Borneo give rise to partially hairy pupae.

Hybridization in the past with a member of the Signifera group (pulchripal-

pis or its ancestor) may be the means by which anopheloides acquired its most

common type of hindtarsal banding. As pointed out in the systematics section

of the Anopheloides subgroup, some larval features may also have been ac-

quired from the Signifera group. Hybridization currently takes place between

anopheloides and albipes and is discussed under that species.

DISTRIBUTION. India, southern China, and southern Japan, south to

Ceylon, Sumatra, Java, Borneo and the Philippine Islands. Material Examined:

636 specimens; 145 0c, 138 9, 1 isolated “ genitalia, 71 P, 281 L; 69 individual

rearings (51 larval, 9 pupal, 9 incomplete).

CEYLON. Aluwilake, 14-26 Nov 1919, R. Senior-White, 10°, 19 [BMNH].

Peradeniya, May-July 1909, 1 0, 39 [BMNH]; K.M. McGahey, 1? [BMNH].

Suduganga Estate, 26 Nov-4 Dec 1919, R. Senior-White, 2 0, 12 [BMNH]; 19

May 1923, R. Senior-White, 2 ¢ [BMNH].

CHINA. Chekiang: Hangchow, 12 Sept 1935, Shih-Cheng Wu, Looe

BMNH|}.

i ok ee OF MALAYSIA. Perak: Pulau Pangkor Laut, 20 Oct 1903,

C.W. Daniels, @ lectotype of nigritarsis (76) [BMNH].

Sabah: Jesselton, 25 July 1956 (US 7), 7 lpo& (7-1, 20-23, 25, 28), 5 lp?

(7-2, 3, 24, 26,27), 1 po (7-29), 11c% (7-30) [BMNH]. Keningau, 11 Aug 1956

(US 55), 3 2 [BMNH].

Selangor: Ulu Gombak, 13 Nov 1956 (096), 1 1p? (096-39) [LIVER]; same

data (0103), 1 lpo& (0103-29) [LIVER]; 10 Jan 1957, J.A. Reid (0129), 51lpo

(0129-6, 11, 13-15), 2 lp? (0129-19, 46), 19 [LIVER], 2 lpo& (0129-44, 45),

2 1p? (0129-25, 42), 1 lp (0129- 44) emul 15 Jan 1957, J. A. Reid (0133),

21po (0133-2,9), 2 1p? (0133-1, 3) [LIVER], 1 o% [USNM]; 14 Feb 1957 (0175),

19 [LIVER]; 13 May 1957 (0197), 1 lpo& (0197-2) [LIVER].

No locality: 18-19 Jan 1922, 1 9 [BMNH].

INDIA. Andaman and Nicobar Islands: Andaman Islands, 22 July 1908,

Lowis and R. White, ? lectotype of maculipes, 3 9 [BMNH].

Assam: Doom Dooma, Sookerating, 11 May 1943, D.E. Hardy (S 31), 3 L

[USNM]; 9 July 1943, G. F. Johnstone (S 31), 1 L [USNM]. Golaghat, 16 May

1925, P.J. Barraud, 1 1p? (1938) [BMNH]. Nongpoh, July 1922, P.J. Bar-

152 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

raud, 1 L [BMNH].

Himachal Pradesh: Simla, 28 Sept 1930, P.J. Barraud, 2 lpo (2541, 2543),

1 1p? (2546), 110 (2542) [BMNH]; S.R. Christophers, 1 0, 19 [BMNH]. So-

lon, Krol Mountain, Aug 1923, P.J. Barraud, 21? (1602, 1603) [BMNH]; 4 July

1930, P. J. Barraud, 2L [BMNH].

Mysore: Voltaour: 1 Oct 1921, P.J. Barraud, 1 1p (916), 4 L [BMNH].

Uttar Pradesh: Dehra Dun, G. M. Giles, © lectotype of anopheloides, 1 2

[BMNH].

INDONESIA. Java: Bogor, 1930, R.W. Paine, 1%, 19, 2P, 5L [BMNH].

Sumatra: Danau Ranau, 2P [UCLA]. Tjoeroek, 4 o, ¢ [UCLA].

JAPAN. Honshu: Kyoto, 1947-1950, p? lectotype of nipponica, 1 po, 6c,

29, 14L[USNM], 1o, 19, 3L [UCLA], 20, 1 L [BMNH], 2L [UCD]. Tok-

yo, 13 July-22 Aug 1949, Lo, 12 (CU. No locality, 6 Oct 1948, 2L [USNM].

NEPAL. Hetaura, June 1955, S.C. Desson, 1 L [BMNH].

PAKISTAN. East Pakistan: Rangamatti, Sept 1922, P.J. Barraud, 1 L

[BMNH].

REPUBLIC OF THE PHILIPPINES. Laguna: Llavac, 14 Sept 1940, P. Suni-

co (TH 111), 1o [BISHOP]. Mt. Makiling, 8 June 1964 (IH 35), 2 L [UCLA]; 1

Aug 1964, (IH 77), 1 L [UCLA]. |

Leyte: Tacloban, 3 Sept 1945, R.E. Gossington, 2 L [USNM].

Mountain: Baguio, 25 Aug 1945, S. E. Shields (443), 219 (443-1, 3) [USNM].

Negros Occidental: Fabrica, Feb 1931, W.V. King (139a), 70°, 29, 5L

[USNM].

Negros Oriental: Cuemos de Negros, 14-16 July 1964, M. Delfinado, 1,

1? ;USNM].

Nueva Ecija: Munoz, 11 Aug 1945, MacMillan, 2 L [USNM].

Occidental Mindoro: San Jose, pee May 1945, E.S. Ross, 1 lp (C-870),

1 1p? (C-55), 60, 49, 16 L[USNM], 1c, 19, 1 L [BISHOP].

Palawan: Irahuan River area, 2 June 1945, D.R. Johnson and J.L. Laf-

foon ou 1 lpo& (851.4) [USNM]; 4 June 1945, J.L. Laffoon (860), 1 lpo&

(860.4), 1 lp? (860.1) [USNM]; 11 June 1945, J.L. Laffoon and Fitzgerald

(875A), 1L [USNM]. Puerto Princesa, 16 Sept 1945 (P 291), 2 1lpo" (291-1, 2),

1 lp, 2L [USNM]. Tarumpitao, 7 Jan 1960, L. Quate, 1L [USNM].

Quezon: Casiguran, 5 June 1957, 40°, 1 9 [UCLA].

Zambales: Subic Naval Base, 18 July 1966 (175), 2 1lpo (175-1, 2) [UCLA].

RYUKYU ISLANDS. Iriomote Island: Shirahama, 17 Nov 1951, R.M. Bo-

Hart; 2c, 3 2 UCD.

Ishigaki Island: Mt. Maeshi, Oct 1951, R.M. Bohart, 12 [UCD]. Teri

River, 17 Oct 1951, R.M. Bohart, 3 L [UCD]. Yarabu Peninsula, Oct-Nov

1951, R.M. Bohart, 8c, 14 9, 9L [UCD], 10, 2 9, 3 L [USNM], 1¢, 19,

3L [CAS], 20°, 29 [BISHOP].

Okinawa: ADA Village, 20 Apr 1955, Nakata, 2 L [USNM]. AHA Village,

6 Oct 1955, I. Zaldua, 1 L [USNM].

SINGAPORE. D.H.C. Given, 30, 29, 1 P, 7 L [BMNH].

TAIWAN. Chu Kod Fan Iu, Chid-I Hsien, 11 Mar 1954, H.H. Chen, 1 o&

[USNM]. Chukou, Fanlu, Chiai, 14 Mar 1954, J.C. Lien (78065), 1 9 [BMNH].

Chung Shan Park, Chiai Chiai, 6 July 1953, J.C. Lien (78020), 1 2 [BMNH].

Chy-Lia, Cu-Ku, Nan-Tou Hsien, 2 Oct 1954, C.L. Chung and J. Y. Li (80662),

1L [BMNH]. Ho-Ping, Tai-Chung, 23 June 1959, J. F. Chiu and W.L. Chen

(00389), 5L [USNM], 4L [BISHOP]. Kechuang, Fanlui, Chiai, 14 June 1955, J.C.

Lien (78008), 1? [BMNH]. Ku Lou, Lail, Fing-Tung, 29 Sept 1958, C.J. Kuo

and C.C. Kang (00047), 29 [USNM]. Mei-Shan, Chia-I, 11 Mar 1954, H.H.

Zavortink: Genus Orthopodomyia 153

Chen and T.S. Lu (75252), 19 [USNM]. Shui-Li, Nan-Tou, 4 July 1954 (80471),

1c’, 29 [USNM]. Sun Moon Lake, 28 May 1948, 39 [BISHOP]. Tinlin, Chushan

Nantuu, 3 Oct 1954, C.L. Chung (80700), 1 L [BMNH]. No locality or date,

J.C. Lien, 3 0° [BMNH].

THAILAND. Chanthaburi: Ban Laem Sing, 1965 (00669), 4 po’ (00669-102,

105, 106,109), 2 p? (00669-107, 114), 29, 2 L [USNM].

Chiang Mai: Doi Pui Huay, 1962 (T 1147), 1%, 29 [USNM]; same data

(T 1148), 30°, 22 [USNM]. Doi Suthep, 25 Oct 1952, D.C. Thurman (M 320),

1 L [USNM]; 30 Dec 1952, D.C. Thurman (M 398), L holotype of lemmonae,

2 L [USNM]; 11 Jan 1953, D.C. Thurman (M 409), 2 L [USNM]; 4 Feb 1953,

D.C. Thurman (M 449), TL [USNM]; 5 Feb 1953, D.C. and E.B. Thurman

(M 462), 2 o [USNM]; a Feb 1953, D.C. and E. B. Thurman (M 545), 1 2 [USNM];

26 Mar 1953, D.C. Thurman (M 638), a0, 29,32; alu enna 3 Dec 1957,

E.B. Thurman (M 856), 1 & genitalia [USNM]; 1962 (T 1086), 19 [USNM];

same data (T 1090), 19 [USNM]; same data (T 1101), 3 9 a eee same data

(T1112), 10° [USNM]; same data (T1113), 10°, 3 9 [USNM]; same data (T 1224),

1 o, 19 [USNM]; same data (T 1256), 1 o& [USNM]; same data (T 1384), 1¢

[USNM]; same data (T 1659), 30°, 22 [USNM]; same data (T 1908), 10, 19

[USNM]; same data (T 1914), 1 2 [USNM]; 1963 (T 1934), 3 o [USNM]; same

data (T 1936), 1 9 [USNM]; same data (T 2061), 1%, 69 [USNM].

Khon Kaen: Tham Pui Tiyan, 1962 (T 1823), 3o%, 19 [USNM]; same data

(T 1824), 19 [USNM].

Nakhon Nayok: Muang, 14 Sept 1963 (NY 37), 1 L[USNM]. Sariga Village,

22 May 1964 (NY 140), 1 1lpo (140-46), 1 1p (140-40) [USNM].

Nakhon Ratchasima: Khao Yai, 1962 (T 1862), 20°, 1 2 [USNM]; same data

(T 1863), 1 o [USNM].

Trang: Kantang, 1964.(TG 133), 1 lpo (133-50) [USNM]. Muang, 1964 (TG

59), 1 lpo’ (59-41) [USNM]. |

No locality or date: (00097), 1 L [USNM]; (00159), 2 L [USNM]; (00303),

11L [USNM]; (00352), 1L [USNM]; (00663), 1L [USNM]; (06671), 2L [USNM];

(T 1045), 1 o [USNM]; (T 1092), 1 o [USNM]; (T 1157), 1 o [USNM]; (T 1251),

6 L [USNM]; (T 1303), 1¢, 19, 1 L[USNM]; (T 1967), 19 [USNM]; (T 1984),

1 9 [USNM]; (T 2084), 10, 12 [USNM]; (T 2087), 19 [USNM]; (T 2573), 1 L

[USNM]; (T 2781), 2 L [USNM]; (CM 33), 1 L [USNM]; (CM 36), 5 L [USNM];

(CM 46), 1 1p2 (46-1-25), 3 L [USNM]; (CM 49), 4.L [USNM]; (CM 114), 2 L

[USNM]; (CM 117), 10° [USNM]; (CM 129), 2L ener (CM 169), 1 lpo’ (169-

2-18), 3 L [USNM]; (CM 171), 1 L [USNM]; (CM 204), 4 L [USNM]; (CM 215),

Lipo’ (215-1-32), 4L a (CM 220), 1 lpo (220-45) [USNM]; (CM 223),

9 L [USNM]; (CM 228), 1 L [USNM]; (CM 229), 1 L [USNM]; (NR 423), 1 L

[USNM]; (PR 167), m3 [USNM]; (RN 36), 1¢ nee

NO LOCALITY. 17 July 1964 (P-37), 2 L [USNM].

NO DATA. 56, 49 [USNM].

Additional records from the literature. CHINA. Hunan: Changsha, Jan-

Mar 1956, L (Chang, 1957a:214, 220, 221, 222).

Kwangtung: Hainan (Chu, 1957: 149, 163).

Yunnan: Chefang, Apr 1940-Mar 1942, L (Chow, 1949:128).

INDIA. Bihar: Sukna, Sept 1922, P.J. Barraud (Barraud, 1927:527-529).

Kerala: Maddathorai, 17 Nov 1908, Annandale, A (Theobald, 1910a:29-30).

Maharashtra: Bhorghat Pass, Apr 1953-Mar 1955, A (Ramachandra Rao

and Rajagopalan, 1957:13, 33).

Punjab: Koti, near Kasauli, Aug 1923, P.J. Barraud (Barraud, 1927:527-

529).

154 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

REPUBLIC OF THE PHILIPPINES. Bulacan: San Jose, Tungkong Manga,

Dec 1929, F.E. Baisas, L,A (Baisas, 1946:35-39).

THAILAND. Nakhon Si Thammarat: Khao Ram, 23 Feb 1922, Pendlebury,

A (Edwards, 1928:53, 57).

NOMEN DUBIUM

24a. Orthopodomyia mcgregori (Banks)

1909. Kerteszia mcgregori Banks, 1909:548-550. TYPE: holotype 0, Basilan

Island, [Zamboanga del Sur], Philippine Islands, 28 Dec 1906, R.C. Mc

Gregor. Holotype was #6666 in the Entomological Collection, ©

Bureau of Science, Manila, P.I., but according to Baisas (1946:34) it was

destroyed during the liberation of Manila in World War II. Reduced to

Synonymy with albipes by Edwards (1932:108); judged to probably be dis-

tinct from albipes by Baisas (1946:34, 35); resurrected from synonymy

with albipes by Knight and Chamberlain (1948:10); reduced to status of

nomen dubium here.

Orthopodomyia mcgregori of Baisas (1946:34, 35).

Orthopodomyia albipes of Edwards (1932:108, in part); Barraud (1934:102, in

part); Bohart (1945:37).

SYSTEMATICS. Kerteszia mcgregori was described by Banks in 1909 from

2 males collected on Basilan Island. The type and presumably the other male

were destroyed during the Second World War. Edwards (1932:108) and Barraud

(1934:102) considered mcgregori to be a synonym of albipes on the basis of

hindtarsal segments 3-5 being white-scaled. Baisas (1946:34, 35) felt that mc-

gregori was probably distinct from albipes and was distinct from the 2 species,

manganus (=anopheloides) and madrensis, which he described from Luzon. He

suggested leaving the taxonomic status of mcgregori uncertain until additional

material was collected from the type locality. Knight and Chamberlain (1948:

10) and Knight and Mattingly (1950:13-15) applied the name to a single male

from Palawan which had the last 3 hindtarsal Segments all white. This speci-

men, however, is anopheloides.

On the basis of hindtarsal ae 3-5 being entirely white-scaled, mc-

gregori could be albipes, andamanensis, anopheloides, madrensis, papuensis,

Siamensis or a distinct Species. The original description indicates that hind-

tarsal segment 2 is white at the apex, implying that it is dark at the base. Al-

So, white spots are mentioned to occur on the bases of wing veins I(R) and V

(Cu), but none is mentioned for the anal vein. Since albipes has white scaling

at the base of hindtarsal segment 2 and always has white scales at the base of

vein 1A, these specimens were not that: species. On geographical grounds it is

unlikely that they were siamensis.

Since it is impossible to tell whether mcgregori is a distinct species or not,

I think the suggestion of Baisas to leave its taxonomic status uncertain until to-

potypic material is available is the best procedure and the name is accordingly

considered here to be a nomen dubium.

DISTRIBUTION. Type from Basilan Island, Zamboanga del Sur, Republic

of the Philippines. Specimens Examined: none.

Zavortink: Genus Orthopodomyia 155

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OMA KDorPwWNF

Zavortink: Genus Orthopodomyia

FIGURES

Distribution of Orthopodomyia

Bancroftia and Folicolae Sections, adult thoracic ornamentation

Orthopodomyia waverleyi, male genitalia and pupa

Orthopodomyia waverleyi, larva

Orthopodomyia signifera, male genitalia and pupa

Orthopodomyia signifera, larva

Orthopodomyia alba, male genitalia and pupa

Orthopodomyia alba, larva

Orthopodomyia pulchripalpis, male genitalia and pupa

. Orthopodomyia pulchripalpis, larva |

. Orthopodomyia species 5, larva

. Orthopodomyia kummi, male genitalia and pupa

. Orthopodomyia kummi, larva

. Orthopodomyia albicosta, male genitalia and pupa

. Orthopodomyia albicosta, larva

. Orthopodomyia fascipes, male genitalia and pupa

. Orthopodomyia fascipes, larva

. Orthopodomyia arboricollis, milloti, sampaioi, vernoni; male genitalia

. Orthopodomyia phyllozoa, male genitalia and pupa

. Orthopodomyia phyllozoa, larva

. Orthopodomyia milloti, larva

. Orthopodomyia arboricollis, larva

. Orthopodomyia flavicosta, male genitalia and pupa

. Orthopodomyia flavicosta, larva

. Orthopodomyia flavithorax, male genitalia and pupa

. Orthopodomyia flavithorax, larva

. Orthopodomyia madrensis, male genitalia and pupa

. Orthopodomyia madrensis, larva

. Orthopodomyia siamensis, male genitalia and pupa

. Orthopodomyia siamensis, larva

. Orthopodomyia wilsoni, male genitalia and pupa

. Orthopodomyia wilsoni, larva

. Orthopodomyia albipes, male genitalia and pupa

. Orthopodomyia albipes, larva

. Orthopodomyia papuensis, male genitalia and pupa

. Orthopodomyia papuensis, larva

. Orthopodomyia andamanensis, male genitalia and pupa

. Orthopodomyia andamanensis, larva

. Orthopodomyia anopheloides, male and female

. Orthopodomyia anopheloides, male genitalia and pupa

. Orthopodomyia anopheloides, larva

177

Fig. |

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F EQUAL AREA PROJECTION

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India

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ORTHOPODOMYIA

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anopheloides

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India

Zavortink: Genus Orthopodomyia

219

INDEX TO SCIENTIFIC NAMES

Aedes 53

Aesculus 8

alba (Orthopodomyia) 7, 8, 9, 11, 12,

18, 26, 27, 28, 29 & 30 (keys), 31,

34, 40, 41, 47, 50, 51

albicosta (Bancroftia, Orthopodomyia)

4.12) °44, 16: 48 Ba 95) oa 64.

66, 71, 80, 90, 95 |

Albicosta group 6, 12, 13, 18, 19&

20 & 21 (keys), 23, 66

*albionensis (Orthopodomyia) 52, 53

albipes (Finlaya, Orthopodomyia) 4, 9,

12, 13;. 14)..48, 72, 93, 209, -170,

111, 112 & 113 & 115 (keys), 119,

126, 182, 133; 1394/9136; 197/°138:

146, 147, 149, 150, 151, 154

Albipes group 6, 7,.12, 13,.18, 19 &

21 & 22 (keys), 23, 56, 71, .89,. 93,

95,107, 108, 5140, 206, 118,181,

£22,125, 129) 433, 136,.138,/439,

140, 141, 149, 150

Albipes subgroup 12, 111, 133, 138

andamanensis (Orthopodomyia) 9, 11,

£2,°18,-93, 110,011) 122 & 1

114 & 115 (keys), 126, 129, 138,

139, 140, 141, 143, 144, 145, 146,

147, 148, 149, 151, 154

Anopheles 9

anopheloides (Finlaya, Mansonia, Or-

thopodomyia) 4, 11, 12, 13, 110,

111, 112 & 113 & 114 & 115 (keys),

118,121) 125,:19ey;490,. 1937 196;

138, 140, 141, 143, 144, 145, 146,

147, 149, 150, 151, 152, 154

Anopheloides subgroup 5, 11, 12, 23,

107, 110, 111,126, 129, 136, 137,

139, 141, 143, 151

arboricollis (Culex, Newsteadina, Or-

thopodomyia) 4, 5, 7, 14, 18, 21,

93, 95, 102, 106, 107, 108, 110,

111, 145

Arboricollis group 12, 18, 20 & 21

(keys), 102

Arboricollis subgroup 5, 107

*bacigalupoi (Orthopodomyia) 75, 76,

*Bancroftia 4, 5, 14

Bancroftia section 12, 14, 16, 19 (keys),

23, 28

*californica (Orthopodomyia) 27, 35,

37, 40, 41, 44, 60

canaria (Serinus) 9

Casuarina 8

Columba 9

Culex 4

Culiseta 18

domesticus (Passer) 9

fascipes (Bancroftia, Mansonia, Ortho-

podomyia, Thomasina) 5, 12, 17,

18, 71, 74, 75 (keys), 76, 79, 80,

82, 83, 84, 90, 107

Fascipes subgroup 5

Ficalbia 18

flavicosta (Orthopodomyia) 11, 12, 13,

14, 16, 91, 92, 107, 108, Od, 110,

111, 112 & 113 & 114 & 115 (keys),

198; (PO) 120) 191, 102,195) tae,

149

Flavicosta subgroup 12, 111, 115, 119,

192,9129) 186

flavithorax (Orthopodomyia) 7, 11, 12,

13, 14, 16, 17, 18, 21 (keys), 91, 92,

107, 108, 109, 110, 111, 112 & 113

& 114 (keys), 115, 116, 118, 119,

120, 122, 125, 129, 149

Folicolae section 12, 14, 18, 19 & 20

& 21 & 22 (keys), 74, 84

fraseri (Culiseta) 18

fuscus (Pipilo) 9

geberti (Orthopodomyia) 3, 94, 95, 97,

100

Grabhamia 4

Group A (Orthopodomyia) 5, 74, 89, 107

Group B (Bancroftia) 5, 89

hippocastanum (Aesculus) 8

Kerteszia 4

kummi (Orthopodomyia) 8, 9, 11, 12,

13,°18,°23, 26, 27, 28 & 20°@ 30

(keys), 34, 40, 41, 42, 56, 58, 60,

63, 64, 65, 80, 90 fe

220

Kummi subgroup 14, 22, 24, 25, 28

(keys), 51, 58

*lemmonae (Orthopodomyia) 110, 147,

150, 151, 153

livia (Columba) 9

*longipalpis (Mansonia, Thomasina) 4,

5, 14, 75, 76, 79, 80

*maculata (Orthopodomyia) 110, 146,

147, 149

*maculipes (Orthopodomyia), 110, 129,

146, 147, 150, 151

madrensis (Orthopodomyia) 12, 71, 92,

108, 109, 110, 111, 112 & 113 & 114

& 115 (keys), 116, 119, 122, 125,

126, 129, 136, 149, 154

*manganus (Orthopodomyia) 146, 149,

150, 154

Mansonia 4, 5, 18

maverleyi (Orthopodomyia) 34

mcgregori (Kerteszia, Orthopodomyia)

3, 110, 147, 151, 154

milloti (Orthopodomyia) 12, 18, 21, 94,

95 (keys), 97, 99, 100, 107, 108

Newsteadina 4, 5, 14

*nigritarsis (Orthopodomyia) 146, 149,

151

*nipponica (Orthopodomyia) 146, 147,

149, 150, 152

nkolbissonensis (Orthopodomyia) 3, 12,

21, 100, 101, 102, 107, 108

Nkolbissonensis group 6, 12, 18, 19 &

20 (keys), 100

Orthopodomyia 2, 4, 5, 6, 7, 8, 9, 10,

bh, 12, in, 19, 19, 41, 10, 74, 79,

89, 95, 102, 106, 107, 108, 126,

136, 149, 150

Orthopodomyia section12, 14, 19 (keys),

91, 93, 95

papuensis (Orthopodomyia) 12, 110, 111,

112 & 113 & 114 & 115 (keys), 129,

133, 138, 139, 140, 141, 142, 154

Passer 9 |

phyllozoa (Bancroftia, Mansonia, Ortho-

wodomyia) 3,7, £2, 84.0107, (28, T1,

80, 84, 85, 89, 90, 107, 111, 133

Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968

Pipilo 9

*Pneumaculex 4, 5, 14

Polystictus 132

Populus 8, 10

pulchripalpis (Culex, Orthopodomyia)

a, O12) 33, 15,22; 25, 2ay 25,

26, 27, 28 & 29 & 30 (keys), 34, 40,

52, 53, 56, 57, 64, 93, 139, 151

Pulchripalpis subgroup 28 (keys), 51,

Quercus 8

Salix 8, 10

-sampaioi (Orthopodomyia) 12, 21, 74,

75 (keys),

Serinus 9

siamensis (Orthopodomyia) 12, 110,

111, 112 & 113 & 114 & 115 (keys),

£19, 121, 126;.129, 130, 193,134,

138, 139, 140, 154

signifera (Aedes, Bancroftia, Culex,

Mansonia, Orthopodomyia, Pneuma-

culex, Stegomyia) 4, 8, 9, 11, 12,

£8o 14) 98, 24, (25, .26,..07,-28, 20

& 30 (keys), 31, 32, 34, 35, 36, 37,

40, 41, 42, 45, 50, 51, 56, 60, 63,

64 :

Signifera group 6, 10, 12, 13, 14, 15,

18, 19 & 20& 21 & 22 (keys), 23,

24, 27, 26: 8, 34, 2, 44, 8 aR

58, 63, 64, 70, 71, 74, 95, 107,

411, 139, 151

Sicnifera subgroup 24, 28 (keys), 30

species 5 (Orthopodomyia) 12:20; 21.

27, 28 & 29 & 30 (keys), 58

species 14 (Orthopodomyia) 12, 19, 20,

21, 102

79, 80, 82, 84, 93

*Theobaldia 18

*Thomasina 4, 5, 14

TFhomasina section 6, 12, 14, 16, 18,

19 & 20 & 2t & 22 (keys), 23, 71,

74, 84, 93, 95, 107, 108, 111

*townsendi (Orthopodomyia) 75, 76, 79,

Toxorhynchites 9

Ulmus 8

Zavortink: Genus Orthopodomyia 221

vernoni (Orthopodomyia) 12, 21, 94 wilsoni (Orthopodomyia) 12, 89, 110,

95 (keys), 96, 97, 99, 100, 107, 111, 112 & 113 & 115 (keys), 119,

110 126, 290, bed, E39, 186; 137

Vernoni group 6,,12, 18, 20 & 22 Wilsoni subgroup 111, 130

(keys), 23, 71, 74, 93, 94, 95, 97,

107,-108, 211 xanthopus (Polystictus) 132

waverleyi (Mansonia, Orthopodomyia,

Pneumaculex) 8, 12, 24, 25, 27, 28

& 29 & 30 (keys), 31, 32, 34, 35,

40, 41, 47, 51, 56, 64

Contributions

of the

American Entomological Institute

Volume 3, number 3, 1968

HEAD MUSCULATURE OF SPHINX MOTHS

(LEPIDOPTERA: SPHINGIDAE)

By Richard C. Fleming