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Contributions

of the

American Entomological Institute

\! S

Volume 8, 1972

CONTENTS

No.1. Gangwere, S. K., M. Morales Martin, and E. Morales

Agacino. The distribution of the Orthopteroidea in Teneriffe,

Canary Islands, Spain. May 5, 1972. 40 pages.

No.2. Arnell, J. Hal and Lewis T. Nielsen. Mosquito studies

(Diptera, Culicidae) XXVII. The varipalpus group of Aedes

(Ochlerotatus). May 9, 1972. 48 pages.

No.3. Zavortink, Thomas J. Mosquito studies (Diptera, Culicidae)

XXVIII. The New World species formerly placed in Aedes

(Finlaya). June 30, 1972. 20 6 pages.

No.4. Kistner, David H. A revision of the termitophilus tribe

Feldini (Coleoptera, Staphylinidae) with a numerical analysis of

the relationships of the species and genera. August 1, 1972.

pages 1-35.

Baker, John N. and David H. Kistner. The reorganization

of the Dorylaerus Complex by the use of numerical analysis

with some new distribution records and new data on their be-

havior. August 1, 1972. pages 36-52.

No.5. Haas, Glenn E., Nixon Wilson, and P. Quentin Tomich.

Ectoparasites of the Hawaiian Islands. I. Siphonaptera.

September 7, 1972. 76 pages.

No. 6. Sirivanakarn, Sunthorn. Contributions to the mosquito fauna

of southeast Asia XIII. The genus Culex, subgenus Eumelano-

myia Theobald in southeast Asia and adjacent areas. October 27,

197 2. 86 pages.

This is an irregular-appearing series,

published by the

AMERICAN ENTOMOLOGICAL INSTITUTE

9950 Warren Road

Ann Arbor, Michigan 48105, USA

Volumes 1-7 available in individual numbers

Volumes 8 and following supplied only as

complete cloth-bound volumes

Contributions

of the

American Entomological Institute

Volume 8, Number 1, 1972

THE DISTRIBUTION OF THE ORTHOPTEROIDEA IN

TENERIFE, CANARY ISLANDS, SPAIN

By

S. K. Gangwere, M. Morales Martin,

and E. Morales Agacino

oe ps -

ae aes ng eye

> TT

THE DISTRIBUTION OF THE ORTHOPTEROIDEA IN

TENERIFE, CANARY ISLANDS, SPAIN?

By

S. K. Gangwere, M. Morales Martin, and E. Morales Agacino”

During the summer of 1970 one of the authors, S. K. Gangwere, carried

out an investigation of feeding in the Orthopteroidea of Tenerife, Canary Is-

lands, Spain, and thereby obtained some knowledge of that island and of the dis-

tribution of certain of its species. Then during his return to the United States

he visited the Instituto Espanol de Entomologia, Madrid, a museum with the

world's largest collection of Canarian Orthopteroidea. These activities yielded

considerable data, and placed him in contact withthe second author, M. Morales

Martin, who for years has been researching the Canarian orthopteroids, has an

extensive private collection, and as an official of the Laboratorio de Zoologia,

Museo de Ciencias Naturales, Santa Cruz, has access to the collections of that

organization. These circumstances presented a unique opportunity for a bio-

geographic study, andthe two entomologists, together withthe third, E. Morales

Agacino, an orthopterist with especial interest in Iberian, North African, and

Canarian species, availed themselves of it to collaborate in the following re-

port, based on the several collections listed above.

During the last fifty years many publications have appeared dealing, in

part, with the Tenerifean orthopteroids. Notable among them are papers by

Chopard (1942, 1946, 1954), Gardner (1960), Johnsen (1970), Willemse (1936,

1949, 1950), Willemse and Bruijning (1949), and Uvarov (1948), which provide

an incomplete faunal list and scattered notes on habits and distribution. Re-

cently, Holzapfel (1970) discussed in detail the systematics and distribution of

the Canarian Acridoidea, and contributed much to an understanding of the bio-

geography of that superfamily. However, there is still neither an up-to-date

list of the entire Tenerifean fauna nor a detailed presentation of the species'

distribution and habits. The present study should help to eliminate that gap

in the literature.

GEOGRAPHIC AND PHYSICAL FEATURES. The Canarian Archipelago

is comprised of thirteen subtropical volcanic islands, seven of which are in-

habited, lying off the Atlantic coast of Africa at a point approximately 195 kilo-

meters from Cape Juby (Riode Oro), 1, 111 kilometers from Gibraltar, andclose

to the Tropic of Cancer. Tenerife, the largest of the included islands, has an

area of 2,053 square kilometers. It is a triangular-shaped body of land whose

hohis project was supported by Grant No. 5560, Penrose Fund, Ameri-

can Philosophical Society, and by a Faculty Research Award from the Gradu-

ate Division of Wayne State University. These grants were for research by the

first author during the summer of 1970.

2Gangwere is Professor, Department of Biology, Wayne State University,

Detroit; Morales Martin, Profesor Mercantil, Poeta Tomas Morales 17, Santa

Cruz; and Morales Agacino,Chief, Departamento de Protecciona la Naturaleza

y al Medio Ambiente, Instituto Nacional de Investigaciones Agronomicas, Madrid.

2 Contrib. Amer. Ent. Inst., vol. 8, no. 1, 1972

vertices are the northeastern Punta de Anaga, the southern Punta de Rasca, and

the northwestern Punta de Teno. The coasts are narrow, often in the form of

rocky, precipitous cliffs, and seldom with beaches or natural ports. The sur-

face is coursed by a central cordillera that extends like a backbone from Anaga

to Teno, and is responsible for relief unusual in an area so small. The island

rises abruptly from the sea, reaches 1, 000 meters above sea level only a few

kilometers inland, and exceeds 2,000 meters at Las Canadas near the center

of the island. Las Canadas is a giant crater from which arises, in turn, the

newer volcano Pico de Teide, whose cone thrusts 3, 716 meters above sea level

into the sky. (See Plate V). 3

The Canaries presumably arose from the sea during the Miocene, and

from that time to the present Tenerife has been subjected to repeated volcanism,

most recently during the 1909 eruptions of Volcan Chinyero. The soil is thin,

even in the woods, where it seldom extends more than 50 centimeters deep. It

has developed fromthe superposition of various volcanic layers of different age,

modified by weathering and erosion, and cut here and there by ravines, or bar-

vyancos, of varying depth. The barrancos are authentic geologic fissures that

act as temporary streams during the rainy season. There are practically no

creeks, other regular water courses, or natural wells on the island; hence, the |

inhabitants rely on artificial water tunnels excavated from the mountainsides

for drinking and irrigation.

CLIMATE. The Canarian Archipelago is close to the Tropic of Cancer,

and the shores are washed by the Gulf Stream and blown by the northeasterly

trade winds, or Alistos. Consequently, the climate is subtropical in nature,

and characterized by minimal seasonal and diel fluctuations in temperature,

humidity, and wind. The temperature of the surface waters of the sea inshore

is approximately 22 degrees C. in summer and 18 degrees C. in winter; and on

land near sea level the lowest mean monthly temperature (January) is 15.3 de-

grees C., while the highest (August) is only 22.4 degrees C. The mean month-

ly relative humidity is remarkably constant, ranging from 53% to 66% at Santa

Cruz and averaging higher inthe north. However, the conditions at particular

stations vary according to exposure and elevation. The general conditions range

from hot and dry at sea level, to rain forest at moderate windward elevations,

to alpine on the high mountains, and the southern section is drier and more

wind-blown than is the northern. (See Table II).

The central mountain range extending the length of the island has already

been mentioned. From easttowest, respectively, it arises abruptly at the Ana-

ga Peninsula; quickly reaches elevations in excess of 1,000 meters above sea

level at points in the Mercedes Mountains; declines below 600 meters of eleva-

tion in the Laguna area; rises again at the Esperanza Mountains; continues with

increasing peaks that terminate eventually at lofty Teide; and then falls at Teno.

This orography has important consequences with respect to the cloud-carrying

winds approaching the island from the northeast. The cloud belt ranges from

approximately 800 meters of altitude to 2,000 or more meters during the win-

ter, but only about 600 to 1, 200 meters during summer. Therefore, the clouds

readily clear the island when in the high, winter configuration, but in summer

are trapped and concentrated along the northern coast, except at the ''Laguna

Belt,'' where some filter past to the south. As aresult, there are two major

climatologic areas in Tenerife: a lushly vegetated, fertile northern zone of

greater humidity and rainfall that lies buried beneath a thick cloud bank during

much of the summer, and a drier southern zone, with reduced rainfall, and

supporting an arid vegetation.

Tenerifean Orthopteroid Distribution 3

Most days are clear or feature intermittent clouds that pass rapidly over

the island, but the degree of cloudiness varies with season and location. Rains

are seasonal, beginning in late September or October, reaching their peak in

November, and declining gradually from December through March. By April

and May precipitation reaches comparatively low levels, and from June into

September it virtually ceases. Thus, the rainy season coincides with fall and

winter, but the temperatures then prevailing are mild, so the period is one of

luxuriant plant growth and flowering in all regions except the high mountains.

Likewise, the dry season coincides with summer heat, causing a pronounced

browning and desiccation of the vegetation in all except the most mesic environ-

ments.

FLORA. The plant communities developed on Tenerife vary with soil,

exposure, elevation, and other factors. From the periphery of the island in-

ward, Ceballos and Ortuno (1951) listed: 1) sparsely vegetated rocky or sandy

beaches or cliffs, 2) coastal scrub, 3) scattered remains of once-extensive ju-

niper forests, 4) laurel forest and heather, 5) pine forest, 6) subalpine scrub,

and 7) alpine barrens. The more important with respect to the Orthopteroidea

are discussed below. (See Plate V).

The semidesert coastal scrub zone, a region comprising almost half of

Tenerife's area, occurs throughout the periphery of the island, but is best de-

veloped in the south. It is a hot, arid, rock-strewn zone populated by a re-

duced number of xerophilic plants including cacti, agave, aloe, spurges (Eu-

phorbia), and the decumbent ice plant Mesembryanthemum.

In the cloud-belt area along the northern face of the island the coastal

scrub yields to evergreen laurel forest(so-called Laurisilva). This plant com-

munity of lauraceous thickets, and with well-developed undergrowth, was once

extensive at moderate elevations along the entire windward side, but has been

reduced by man's activities, so that the few remaining stands are restricted

to the comparatively isolated Anaga Peninsula. There it intermingles with

arborescent heather (Fayal-brezal), and constitutes a living remnant of the Ter-

tiary rain forests of Europe, now extinct on the continent (Lems, 1960).

At levels above 1, 000 to 1,500 meters of elevation the laurel and heather

- plant communities are supplanted by pine forest (Pinar). These extensive co-

niferous woodlands are of several types, but the dominant species is Pznus

canariensis, a now-indigenous pine that, during the Tertiary Epoch, also grew

in Europe. P. canariensis forms magnificent, thick forests in Tenerife, but

supports an impoverished undergrowth, so the insect fauna associated with it

is reduced.

At approximately 2,000 meters above sea level there appears a subalpine

scrub zone (Retama-codeso), chiefly of the legume shrub Spartocytisus nubi-

genus interspersed with a reduced number of herbs. The few plant species

there occur in widely separated patches that take root in volcanic ash covered

by sand and pumice.

Above approximately 2,700 meters of elevation the subalpine scrub gives

way to austere alpine barrens. Once into the latter, nothing grows except a

lichen, a moss, and Viola cheiranthifolia, found living in the lava up to with-

in a few meters of the very summit of Teide.

The foregoing concerns the natural regions of Tenerife, but many parts of

the island are not natural, being extensively cultivated, and here andthere ur-

banized. The most important crop plant grown is banana, but orange, tobac-

co, tomato, potato, grain, almond, onion, and many other crops are planted.

4 Contrib. Amer. Ent. Inst., vol. 8, no. 1, 1972

Along the roadsides are found eucalyptus, cedar, mimosa, casuarina, palm,

and other trees, as well as a profusion of flowering jasmine, heliotrope, da-

tura, geranium, bougainvillea, magnolia, poinsettia, and other tropical and

subtropical vines, shrubs, and herbs, many of them introduced.

The flora of Tenerife is incredibly extensive and diversified for so small

an area. Ina total land area of only 2, 053 square kilometers are found 1, 079

species of vascular plants, 91 of which are endemic to this island and another

247 endemic to the Canaries and related islands (Lems, 1960). This flora, an

admixture of the native and the introduced, is comprised largely of Mediterra-

nean, endemic, and cosmopolitan species, but also includes Ethiopian and other

elements, mostly of a tropical or subtropical nature.

Approximately one-third of the Canarian plants are endemic. This height-

‘ened incidence of endemism presumably stems from geographic isolation and

distinct ecological zonation and stratification, stemming from an orography- .

induced multiplicity of climates and microclimates--hence, much of the inter-

est that the Archipelago holds for botanists. No less significant is the exist-

ence in the Canaries of certain Tertiary species. These plants, which include

the laurels mentioned earlier, as well as the Canary Pine and the Canary Drag-

on-tree (Dracaena draco), a gnarled tree lily, probably reached the islands

during the Miocene or Pliocene, and constitute a present-day reproduction of

the ancient forests that once prevailed in southern Europe; today they are ex-

tinct on the continent, represented only by fossils (Lems, 1960).

COLLECTION LOCALITIES. One hundred twenty-nine collection locali-

ties are herein cited for Tenerife. They are alphabetized and numbered,

which permits space-saving numerical designation in Table I. The code is as

follows:

1) Adeje; 2) Afur; 3) Agua Garcia; 4) Agua Garcia, Ravelo de; 5) Aguaman-

sa; 6) Aguamansa, Los Organos de; 7) Aguirre, Monte; 8) Anaga, Monte; 9) Ar-

ico; 10) Arafo; 11) Arico Viejo; 12) Arico, Las Montanetas de; 13) Bailadero=

El Bailadero; 14) Bajamar; 15) Bermeja, Montana; 16) Buenavista; 17) Bufadero,

Barranco de=Barranco Bufadero; 18) Canadas=Las Canadas del Teide; 19) Can-

delaria; 20) Catalanes, Alto de; 21) Chio; 22) Cristianos=Los Cristianos; 23)

Cuesta=La Cuesta; 24) Cuesta, Montana Guerra de la; 25) Diego Hernandez,

Canada de; 26) Esperanza=La Esperanza; 27) Esperanza, Hoya de las Raices

de la; 28) Fuente Fria; 29) Fuente Guillen=Hortigal Alto; 30) Fuente Joco; 31)

Galletas=Las Galletas; 32) Garachico; 33) Garachico, La Montaneta, Los Cas-

tanos de; 34) Gigantes=Los Gigantes; 35) Granadilla; 36) Granadilla, Pinar de;

37) Grande, Barranco=Barranco Grande; 38) Guajara, Pie de; 39) Guamasa;

40) Guancheros, Canada de los; 41) Guayonje, Barranco de; 42) Guerra, Valle

de; 43) Guimar; 44) Guimar, Arenales de; 45 Guimar, Montana Grande de;

46) Guimar, Volcan de; 47) Hierro, Barranco del; 48) Hondo, Barranco=Bar-

ranco Hondo; 49) Icod; 50) Icod, Las Abiertas de; 51) Icod, Pinar de; 52) Icod,

Volcan Chinyero de; 53) Igueste; 54) Igueste de S. Andres, Carretera de; 55)

Infierno, Barranco del; 56) Izana; 57) Jardina; 58) Jimenez, Valle de; 59) Jun-

cos, Barranco de los; 60) Laguna=La Laguna; 61) Laguna, Gallardina; 62) La-

guna, San Roque de la; 63) Laguneta; 64) Laguneta Alta; 65) Lagunetas=Las La-

gunetas; 66) Lena, Barranco de la; 67) Llano los Loros; 68) Llano de Maja;

69) Martianez, Barranco de; 70) Masca; 71) Masca, Valle de; 72) Matanza=La

Matanza; 73) Medano=El Medano; 74) Medano, Arenales del; 75) Medano, Mon-

tana Roja del; 76) Medida, Costa de la; 77) Mercedes=Las Mercedes; 78) Mesa

Mota; 79) Mesas=Las Mesas=Monte de las Mesas; 80) Monton de Trigo, Cana-

Tenerifean Orthopteroid Distribution 5

da del; 81) Ninfa, Barranco de la; 82) Ofra, Montana de; 83) Orotava=La Oro-

tava; 84) Poris de Abona; 85) Portillo=El Portillo; 86) Portillo, Canada del;

87) Puerto de la Cruz; 88) Punta del Hidalgo; 89) Rambla=La Rambla; 90) Re-

alejo; 91) Rodeos=Los Rodeos; 92) Roque de Caramujo; 93) San Andres; 94) San

Andres, Barranco de; 95) San Andres, Carretera de; 96) San Diego; 97) San

Isidro; 98) San Isidro, Carretera de; 99) San Jose, Barranco de; 100) San Juan

de la Rambla; 101) San Juan, Puerto de; 102) Santa Cruz; 103) Santa Cruz, Bar-

rio Salud Alto de; 104) Santa Cruz, Finca Oramas de; 105) Santa Ursula; 106)

Santa Ursula, Finca Malpais de; 107) Santiago, Valle de; 108) Santos, Barranco

de=Barranco Santos; 109) Sauzal; 110) Seco, Barranco de=Barranco Seco; 111)

Silos, Monte de los=Monte del Agua; 112) Tabares, Valle de; 113) Taco, Mon-

tana de; 114) Tacoronte; 115) Taganana; 116) Taganana, Vueltas de; 117) Tahodio,

Barranco de=Barranco Tahodio; 118) Tahodio, Charca de; 119) Tanque=El

Tanque; 120) Tegueste; 121) Teide, Alta Vista del; 122) Tejina; 123)Tejina, Cos-

ta de; 124) Ten-Bel, Las Galletas; 125) Teno; 126) Tigaiga, Ladera de; 127) Ve-

ga=La Vega; 128) Vilaflor; 129)Vilaflor, Pinar de.

RESULTS

The results of this study are summarized in the Biological Notes and in

Table I. That table, presented in the final section of this report, gives a pro-

visional list of the Tenerifean species, their world distribution, their distri-

bution within the island, their Tenerifean distribution type, and their season-

al periodism.

DISCUSSION

FAUNA. One would expect the Canarian orthopteroid fauna, an insular

assemblage, to be more impoverished than are the continental faunae at the

same general latitude, and indeed this is the case; the Canaries correspond in

latitude to southern Morocco, where the fauna is undeniably richer. However,

for a fauna isolated on a small archipelago separated by a wide expanse of wa-

ter from others, the Canarian fauna is comparatively rich and diverse. That

of Tenerife, the only Canarian fauna here treated, numbers 88 species (13

Blattoidea, 7 Mantoidea, 12 Tettigonoidea, 15 Grylloidea, 29 Acridoidea, and

12 Dermaptera) belonging to 31 subfamilies and 58 genera. Thus, this fauna

is reasonably representative, for among its species are included members of

all major superfamilies except the Phasmoidea, or walking-sticks. (See Ta-

ble I).

Much of the interest that the Tenerifean fauna has elicited in the past is

based not on numbers of species and groups but on the presence of species with

unusual geographic distribution. Many are autocthonous; a total of 24 of the

species (27%) and 6 of the genera (10%) are endemic either to Tenerife or to

the Canaries in general. The percentage of endemic species per group varies

somewhat, ranging from 58% (Dermaptera), to 33% (Tettigonoidea), to 29%

(Mantoidea), to 23% (Blattoidea), to 21% (Acridoidea), to 13% (Grylloidea). Sev-

eral of the endemics appear to be relicts from faunae that in long-gone ages

inhabited other parts of the world. A good example is Anataelia canariensis,

a representative of the archaic earwig group Protodermaptera (Fernandez,

1966).

The affinities of the Tenerifean fauna are mostly North African, fora

good number of its species are either native to, or closely related to, forms

6 Contrib. Amer. Ent. Inst., vol. 8, no. 1, 1972

of nearby Morocco and related lands. Nevertheless, it also contains many gen-

eral Mediterranean, Ethiopian, and cosmopolitan species. An examination of

Table I reveals that 30 Tenerifean species are Mediterranean or Mediterrane-

an- Palearctic in world distribution; 16 species Ethiopian, Mediterranean and

Ethiopian, or Mediterranean and Afro-Asian; 14 species cosmopolitan or tro-

picopolitan; 4 species essentially Holarctic; and the remaining 24 species en-

demic. With respect to the Tenerifean genera 15 prove to be essentially Med-

iterranean or Mediterranean-Palearctic in world distribution; 2 genera Ethio-

pian; 4 genera occur both within the Ethiopian and the Palearctic realms; 19

genera cosmopolitan or tropicopolitan; ll genera globally widespread, being

Ethiopian, Palearctic, Oriental, and sometimes even Australian or Neotrop-

ical in distribution; and 6 genera endemic.

One-hundred per cent of the autocthonous species of the superfamilies

Blattoidea, Mantoidea, Tettigonoidea, Grylloidea, and the order Dermaptera

prove to be either brachypterous or apterous in at least one of the sexes. This

suggests a direct, positive correlation between endemism and meiopterism

(brachyptery or aptery), a relationship not invalidated by the fact that several

non-endemic species also have reduced wings. However, the correlation is

not invariable, for 6 species of Acridoidea are endemic to Tenerife, yet only

two are brachypterous.

These facts have a bearing on adaptive radiation. Water is a barrier to

orthopteroids and many other terrestrial insect species, which means that a

given island comes to be inhabited only by those forms able to immigrate.

Once a terrestrial population has surmounted this barrier, and has become

established in the isolated environment, it may, comparatively speaking, be

without competition, and quickly undergo adaptive radiation to meet the vary-

ing requirements of particular habitats. In Tenerife and similar islands,

strongly isolated geographically, and also highly zoned and stratified ecologi-

cally, the potential habitats open to species are numerous, and the occupying

fauna may develop in diversity beyond that normally expected.

Based on the foregoing, it is apparent that Tenerife has a comparatively

high percentage of endemic species that, with the exception of the acridoids,

are of reduced vagility owing to meiopterism. Moreover, they tend to be spe-

cialized in habitat selection. Some of them, for example, the earwig Anatae-

lia canariensis and the katydid Canariola nubigena, could be relict forms that

long have inhabited Tenerife, but most are assuredly recent to the island.

Clearly, even the brachypterous and apterous orthopteroids are capable of be-

ing transported passively the approximate 195 kilometers from the African

mainland, and many could have come to occupy the island in that manner,

while the stronger fliers, such as the locust Schistocerca gregaria, can easi-

ly negotiate this distance by wing.

The evolution of distributional patterns, endemism, and their possible re-

lation to meiopterism are of tremendous interest, but require detailed analysis

of orthopteroid distributions on other islands of the archipelago and on the con-

tinent. Inasmuch as the data herein pertain only to Tenerife they are insuffi-

cient to enable anything more ambitious than has already been attempted. How-

ever, the reader may turn to Holzapfel's excellent discussion of these matters

with respect to the Canarian Acridoidea (1970).

DISTRIBUTIONAL ANALYSIS. A provisional list of species was compiled

using the collections of M. Morales Martin, those of the Laboratorio de Zoolo-

gia, and those of the Instituto Espanol de Entomologia. This list, modified in

Tenerifean Orthopteroid Distribution "

accordance with recent literature, was then used as the system against which

all available distributional data were plotted. The locality data of each spe-

cies were plotted on an individual map, the maps compared with one another,

and from these data a classification of distributional types erected. This in-

formation, together with remarks on world distribution and seasonal period-

ism, is presented in Table I.

The basic information just mentioned is, of course, nothing more than a

series of dots on a map, each representing a particular locality from which

one or more specimens of a species have been taken. A hypothetical distri-

bution was derived therefrom by assuming that the range is continuous, which

allows the dots to be interconnected to form an area on the map generally sug-

gestive of the species’ true distribution. For example, if all records are clus-

tered along the northern coast, a northern distribution is assumed. Natural-

ly, the records are incomplete and not necessarily representative, though they

are a product of random collecting by many entomologists over a period of

three quarters of a century. One example is sufficient. Pattern VIIi (Plate

IV) is a distribution in which a species occurs in the southern coastal area ad-

jacent to the west coast but is lacking in the west, a similar area. Failure to

be recorded from the west could stem from the fact that it does not live there,

but is probably a consequence of collecting effort. The western area is remov-

ed from the island's two big population centers, and until recently did not even

have a good road that enabled ready access to it.

An elaborate classification of the biogeographic areas of Tenerife was ad-

vanced by Ceballos and Ortuno (1951), based primarily on plant distribution.

(See Plate V). Therefore, it was desirable to plot the orthopteroid distribu-

tions obtained during the present study against their published scheme. The

fit, or lack thereof, was determined by using the following criteria: 1) the

range hypothesized for a given species is based primarily on the available dis-

tributional records; 2) the range is broadened to include that part of the natu-

ral region in which the records fit, and is terminated at the adjacent physical

and/or climatic barrier; and 3) the range may overlap two or more natural re-

gions, but there can be no gap between.

The result obtained by plotting the orthopteroid distributions against the

Ceballos-Ortuno zones proved less than satisfactory. Approximately one-

third of the orthopteroid species fitted adequately into one or another distribu-

tional patterns consistent with the Ceballos-Ortuno scheme, but one-third did

not fit at all, and another one-third proved intermediate in this respect, nei-

ther violating the scheme nor being entirely consistent with it. This is consis-

tent with Holzapfel's findings (1970); she concluded that it is not feasible to at-

tempt classification of the Canarian acridoids according to distribution in flor-

istic zones.

It was considered that a much better fit might be obtained by plotting the

orthopteroid distributions against elevation as modified by exposure. The fit,

or lack thereof, was investigated using the same criteria as before, and the

results indicate a gratifying overall agreement. Only two species, Canariola

nubigena and Guanchia canariensis, have ranges inconsistent with elevational

zones aS modified by exposure; four other species proved somewhat inconsis-

tent; and the remaining 76 species were consistent with the new scheme.

PATTERNS OF DISTRIBUTION. The orthopteroid distributions examined

fall into 37 provisional patterns (explained in Plates I-IV) that can be aggrega-

ted into seven major categories, as follows:

8 Contrib. Amer. Ent. Inst., vol. 8, no. 1, 1972

I, Ubiquitous or essentially ubiquitous. 8 spp. The insects with thisdis-

tribution are recorded from throughout the island, except sometimes the heights

of Las Canadas or the western coast.

TI, Southern. 8 spp. Two of these distributions are little more than ex-

panded southern coastal distributions, while the third (Pattern Ila) encompas-

ses all but the northern coast of the island.

Ill. Eastern. 5 spp. These distributions ipewiee roughly the eastern

third of the island, except for Pattern IIId, which is a narrow tongue-like dis-

tribution extending from Santa Cruz to Esperanza.

IV. Northern. 14 spp. These insects have ranges that are little more

than expanded northern coastal distributions, except for Pattern IVa, which

takes in the entire north, as well as the central heights of Las Canadas, and

Pattern IVe, which includes the northcentral area, as well as a tongue of land

curling southward around Las Canadas.

V. Eastern crescent. 7 spp. This distribution is characterized by occu-

pancy of both northern and southern coasts as joined inthe east. Such patterns

are but expanded coastal distributions, and their failure to extend onto the west-

ern coast is possibly explicable on the basis of insufficient collecting.

VI. Interior. ll spp. These distributions involve one or another segments

of the central mountain range that extends the length of the island from Anaga

to Teno.

VII. Coastal. 29 spp. These littoral distributions, from coastline up to

about 600 meters above sea level, involve the entire coast, or one or another

parts of it, and sometimes the ''Laguna Belt."’ The latter is a low, inland ex-

tension of coast between the Mercedes and Esperanza Mountains, to the east

and west, respectively. (See Patterns VIId, Vile, and VIIf). The southern and

western coasts of the island are similar, so the absence of certain southern

coastal species from the west (for example, Pattern VIIi) is possibly explica-

ble on the basis of insufficient collecting.

Based on the foregoing we can characterize the distributional patterns of

the major orthopteroid groups as follows:

The Blattoidea have an essentially coastal distribution; only one species,

Arbiblatta bivittata, extends any great distance into the interior.

One mantis, Ameles limbata, is virtually ubiquitous ; two others, A. gra-

cilis and Pseudoyersinia teydeana, have interior distributions; and the remain-

ing species are essentially coastal insects.

Except for the ubiquitous Phaneroptera nana sparsa the Tettigonoidea, or

katydids, are restricted to the north or northeast of the island, especially in-

terior zones.

The interior Gryllomorpha canariensis, the eastern Acheta meridionalis,

the northern Gryllus bimaculatus, and the northern and central Oecanthus pel-

lucens are non-coastal species of crickets. The remaining Grylloidea are es-

sentially coastal in distribution, though some extend well inland.

The distributional patterns of the Acridoidea do not readily lend them-

selves to characterization, for, as a group, they are highly varied. Some

species are essentially ubiquitous; some coastal; some eastern; some north-

ern; and some are interior forms.

The Dermaptera, like the Acridoidea, are variable in their distribution.

Some species are distributed throughout the north or the south of the island;

some in the interior; and some along the coast, often with considerable exten-

sions inland.

Tenerifean Orthopteroid Distribution 9

ORTHOPTEROID FAUNAL REGIONS. According to the Ceballos-Ortuno

scheme (1951), based primarily on plant distribution, the island of Tenerife is

characterized by the following biogeographic regions: low scrub, juniper for-

est (Sabinar), laurel forest (Laurisilva), pine forest (Pinar), scrub (Escobo-

nal), subalpine scrub (Retama-codeso), and alpine barrens (Violeta). (See

Plate V). We have noted that the island's orthopteroid distributions are incon-

sistent with this plan. There is partial agreement, but, of course, that is ex-

pected, for the Ceballos-Ortuno zones are themselves predicated in part on

elevation, climate, and exposure.

Based on the present study, there is an additional zone to be recognized,

the beach zone; and the juniper forest, escobonal scrub, and alpine barrens

are not important in the distribution of orthopteroids; the laurel forest is of

some importance, though it does not support a unique fauna; and the low scrub,

pine forest, and subalpine scrub are unique areas populated by distinct orthop-

teroid faunae.

Let us comment on the important zones, paying especial attention to or-

thopteroids unique and exclusive to them (marked u, below) and those so com-

monly encountered there as to be considered typical of them (¢, below).

The beach zone features a rocky or sandy substrate, black because of the

island's volcanic origin, and a depauperate vegetation. Several orthopteroids

belong here: Acvotylus i. insubricus (t), Anataelia canariensis (u), Labidura

riparia (u), Mogoplistes squamiger (u), Wernerella picteti (t).

The sublitioral scrub zone is best developed along the southern coast at

elevations below 600 meters. It is an arid, semidesert region supporting a

reduced number of xerophilic plants. The orthopteroid fauna that frequents

it is best characterized by: Calliptamus plebeius (t), Oedipoda canariensis

i“ Sphingonotus spp. (t), Stenohippus epacromioides (u), Wernerella picteti

t).

The tropical forest zone, or laurel forest, is an area of lush plant growth

developed in that part of the northeastern face of Tenerife that is exposed to

the summer cloud belt. Here, among the wooded ravines occurs a somewhat

distinctive orthopteroid fauna including: Arbiblatta bivittata (t), Ariagona mar-

garitae (t), Calliphona k8nigi (t), Platycleis spp. (t), Tessellana tessellata (t).

However, several of these elements are shared with the central forest zone,

and none seems to be unique.

The central forest zone, or pine zone, occurs at middle elevations up to

about 2,000 meters above sea level, and supports meadows and other kinds of

communities, as well as luxuriant pine forests. Prominent among the orthop-

teroids that occur here are: Acvotylus patruelis (t), Ameles gracilis (u), Cal-

liptamus plebeius (t), Canariola willemsei (u), Platycleis sabulosa (t), Tes-

sellana tessellata (t).

The subalpine scrub zone is located above 2, 000 meters of elevation. It

is populated by a sparse growth of herbs and legume shrubs separated by ex-

panses of bare ground, and is characterized by: Avrbiblatta bivittata (t), Cal-

liptamus plebeius (t), Pseudoyersinia teydeana (u), Sphingonotus willemsei (u).

The above classification of Tenerifean distributional zones is somewhat

different from that postulated by Holzapfel (1970), who based hers on the occur-

rence of acridoids throughout the entire archipelago. She recognized open

beach, below 300 meter xerophytic, 300-500 meter forest, and above 1,500

meter subalpine vegetational belts. The differences between the two systems

appears mostly discretionary. Whether one accepts 300 meters of elevation,

600 meters, or some figure between those values as the boundary betweenthe

10 Contrib. Amer. Ent. Inst., vol. 8, no. 1, 1972

xerophytic and forest communities depends on a number of variables, including

location (whether on the northern coast, where the lower estimate is preferable,

or on the southern coast, where the higher figure is probably more realistic), as

well as the species of concern. Holzapfel dealt only with the acridoids, where-

as the present report concerns the Orthopteroidea as a whole. Likewise, the

choice between 1,500 meters of elevation or 2,000 meters must be based in good

part on discretion, for that boundary, too, varies with location. Finally, there

is the rain forest that is here recognized, though with reservations, but is ag-

gregated with the general forest category by Holzapfel.

PHENOLOGY. The seasonal occurrence of the Orthopteroidea is as much

an aspect of their distribution as is place of occurrence. Therefore, it is of in-

terest to consider their annual cycle on Tenerife. However, at the outset it

must be stressed that the seasonal data available to the writers (see Table I) are

in some cases fragmentary.

In general, the Orthopteroidea respond to the mild, subtropical Canarian

climate in the expected manner. They tend to be active all year long, and ei-

ther nymphs or adults may be taken at any time from within the preferred habi-

tats. These remarks pertain, of course, to species found below 1,500 meters

of elevation and especially to littoral forms in areas exposed to the sun. In con-

trast, the orthopteroids that are restricted to subalpine and other high zones are

forced into a diapause during the unfavorable winter months, and do not appear

as nymphs until spring and as adults until summer.

Inherent within most orthopteroids is a pronounced population fluctuation

that is only in part taken into account in Table I; all that table shows is that anin-

dividual of a given species was collected during a particular month, but that spe-

cies may actually have been so reduced in number that it was almost non-exis-

tent at that time. Such population fluctuations are maximized in the case of sub-

alpine forms and minimized in littoral forms.

Several patterns of seasonal occurrence are discernible with respect tothe

major orthopteroid groups of Tenerife (Table I):

The Blattoidea are coastal forms, and, in accordance with that subtropi-

cal distribution, can be found adult all year long. The majority of the species

are domiciliary, and in cases where sufficient records are available appear to

produce many generations per year. The native species, too, are adult through-

out the year, and at some points within their range are probably also continuous

reproducers throughout the year.

The Mantoidea have an adult occurrence that encompasses spring and sum-

mer and occasionally fall.

The Tettigonoidea prove more variable than do the foregoing, but all ex-

cept the ubiquitous Phaneroptera nana sparsa have a definite seasonal periodism.

In most instances it is one in which adults occur from spring until autumn.

The Grylloidea tend to be adult throughout the year, a condition in accord-

ance with their mostly coastal distribution. It is difficult to say whether they

produce one or several generations per year, but probably those several spe-

cies that are semi-domestic are of the latter type.

Virtually all of the Acridoidea for which there are sufficient data to make

an assessment can be found in the adult condition throughout the year. The ma-

jor exceptions are Calliptamus plebetus, a ubiquitous species, Schistocerca

gregaria, a plague locust that repopulates the island as the fall winds from the

Sahara descend upon it, Sphingonotus willemsei, a subalpine species, and Oe-

daleus decorus. However, it is doubtful that the acridoids as a group produce

Tenerifean Orthopteroid Distribution ll

more than one generation per year; they are simply long-lived.

In general, the Dermaptera give evidence of being adult throughout the

year, and most species probably have several generations per year. The pos-

sible exceptions are a few species in which the data are too incomplete to per-

mit evaluation.

BIOLOGICAL NOTES

Arbiblatta bivittata. A delicate endemic cockroach, winged in the male

sex, wingless in the female, this species is found throughout the northern part

of Tenerife. It is especially characteristic of the laurel and pine forests, where

it takes refuge in humus, soil crevices, leaf litter (especially of Aeonium), tree

hollows, and under stones. It also lives under similar conditions up to 2,500 me-

ters in the high mountains and less often in the lower zones of the north down to

300 meters above sea level. Adults are abundant all year long throughout the

range, except at Teide during the harsh winter months.

Blattella germanica. A domestic species characteristic of the coastal dis-

tricts, especially the port cities, of Tenerife. Adults occur throughout the year.

Loboptera canariensis. This small, wingless endemic is taken at Santa

Cruz and San Diego, a distribution within the ''Laguna Belt, '' which is an inland

extension of coast between the Esperanza and Mercedes Mountains.

Loboptera decipiens. Under leaves, stones, and debris in woodlands from

Esperanza through into Mercedes.

Loboptera fortunata, Another endemic, this moderate-sized, wingless

cockroach is found along the northern coast and Anaga Peninsula of Tenerife. In

the latter it is readily taken under stones at El Bailadero, but is never truly a-

bundant. It is adult throughout the year.

Blatta orientalis. A domestic species that occurs at El Medano and La La-

guna, a southern coastal distribution.

Periplaneta americana. A domestic species with a coastal distribution in

Tenerife and especially characteristic of the port cities. Adults are taken

throughout the year.

Periplaneta australasiae. A domestic species recorded from the ''Laguna

Belt, '' a coastal distribution.

Periplaneta brunnea. The single specimen taken of this domestic species

is from La Laguna.

Leucophaea maderae. This cosmotropical cockroach, another domestic

Species, has a coastal distribution in Tenerife.

Pycnoscelis surinamensis. A domestic cockroach found in cities and vil-

lages throughout coastal Tenerife. Adults are not recorded from autumn, but

probably occur then too.

12 Contrib. Amer. Ent. Inst., vol. 8, no. 1, 1972

Zetha vestita. This small cockroach, winged in the male sex, vestigial-

winged in the female, is an adventive from the Neotropical Realm. It is here

recorded from boxes of bananas in San Isidro, La Laguna, and Bajamar (anoth-

er version of the ''Laguna Belt'') and from Santa Ursula along the northern

coast. It is adult throughout the year.

Leurolestes circumvagans. A few specimens of this domestic cockroach

are at hand from Granadilla and Santa Cruz, a southern coastal distribution.

Ameles gracilis. This mantis, winged in the male sex, vestigial-winged

in the female, has an inland distribution. It lives on the ground or on shrubs

from the pine forests of Esperanza through Aguamansa to the Canada de Diego

Hernandez. The available records of adults suggest a summer-fall periodism

for that life stage.

Ameles limbata. This thamnophilous mantis, similar to the preceding in

appearance, size, and habits, is more widespread, having an essentially ubi-

quitous distribution within Tenerife. Nonetheless, it is never abundant. Adults

are encountered from April through December.

Pseudoyersinia subaptera. A small vestigial-winged mantis, this endemic

has a southern distribution. It is phytophilous, generally on herbs and small

shrubs.

Pseudoyersinia teydeana. Another vestigial-winged endemic, this mantis

is found above 2,400 meters at Las Canadas del Teide, where it lives on herbs

and shrubs, and deposits its eggs below stones. It is never abundant. Adults

are encountered from March through August.

Mantis religiosa. This mantis is widespread in many parts of the Old and

New Worlds. It is found in several Tenerifean localities, but is absent from

the westand from the central heights about Teide, giving it an essentially cres-

centic distribution. It is most common near Los Rodeos and Ten-Bel, where

adults are encountered on low vegetation from June through November.

Blepharopsis mendica. This large, thamnophilous mantis inhabits dry

zones along the southern and northeastern coasts of Tenerife. Adults are re-

corded from March through October.

Hypsicorypha gracilis. This slender-bodied, phytophilous mantis has an

essentially coastal distribution, with extensions inland across the Anaga Penin-

sula. On the basis of available records, adults seem restricted to spring and

summer.

Phaneroptera nana sparsa. This small, delicate-bodied, fully winged ka-

tydid is widespread throughout the Old World, and is ubiquitous within Tenerife.

It frequents both cultivated and wild situations, and is thamnophilous. Adults

are encountered during all months of the year. .

Canariola nubigena. This endemic meconemine is a small, vestigial-wing-

ed katydid with a caudally tumid pronotum. Based on previously published in-

formation, it is known to occur in the pine forests of Aguamansa (Willemse,

Tenerifean Orthopteroid Distribution 13

1936). However, in recent years it has also proved available in numbers in

the coastal locality of Bajamar and inland in the Mercedes Forest and adjacent

areas, where it lives on herbs and shrubs. Adults are encountered in the

spring from February through June.

Canariola willemsei. This endemic is similar to C. nubigena in habits,

size, and general appearance except for its shiny body surface. It is record-

ed only from Tacoronte and Aguamansa, an inland distribution, and on the ba-

sis of the few available records seems to be mature in autumn.

Calliphona kénigi. This large, powerful predator extends throughout the

north of Tenerife, to which it is endemic. It is characteristic of inland woods

in that part of the island, but occasionally reaches lower zones, such as at Ba-

jamar, Guimar, and San Andres. Nymphs up to 3 cms. in length are easy to

find in the typical localities, where they tend to frequent buttercup flowers.

However, adults are not so often encountered, presumably owing to an arbo-

real habit. Adults have been repeatedly verified for the spring and summer

months, and are also seen infrequently through autumn.

Ariagona margaritae. A small, vestigial-winged endemic that lives inthe

north of the island, especially at the edges of laurel forest above approximate-

ly 700 meters of elevation. Typical localities are Monte de las Mercedes,

Las Lagunetas, and Monte de los Silos. It is found there among herbs, and

is adult from July through November. It is extraordinary for its leaps, which

are of unusually great velocity for a katydid of that size.

Platycleis grisea. A few specimens of this katydid are at hand from Ta-

coronte and La Esperanza, an inland distribution.

Platycleis intermedia. " A few specimens are here listed from the ''La-

guna Belt.'' Adults are recorded from April through July.

Platyclets sabulosa. ‘ This moderate-sized, fully winged katydid has a

northeastern and central distribution on Tenerife, where it is geophilous in

dry cultivated zones or wild zones, especially in the center of the island at an

elevation of approximately 500 to 1,000 meters above sea level. Records of

adults are available from March through October, but the seasonal periodism

could well extend beyond those limits.

Tessellana tessellata. Not unlike a small Platycleis (with which genus it

was grouped for many years), this species has an eastern distribution and an

adult occurrence that extends from March through November.

Decticus albifrons. This large, powerful, fully winged katydid is common

3 The members of this genus are variable, poorly understood, and diffi-

cult taxonomically, so determinations always require great care and compe-

tence. Therefore, the records included herein, as well as those of earlier

authors, should be viewed in that light.

14 Contrib. Amer. Ent. Inst., vol. 8, no. 1, 1972

in cultivated fields and other open places throughout the eastern and central

parts of the island, and is phytophilous. Its adult periodism extends at least

from March through November. This rapacious predator is widespread else-

where, being found throughout the entire Mediterranean region.

Gryllus bimaculatus. A large, dark-bodied cricket with a palish suffusion

at the base of each tegmen, G. bimaculatus is the most common gryllid in the

Canary Islands, and elsewhere occurs through the whole of Africa and in ma-

jor parts of Europe and Asia. It occurs throughout Tenerife, but is most com-

mon in the coastal zones from Puerto de San Juan through Pinar de Icod to Gui-

mar and beyond through the Anaga Peninsula. It is found on the ground in cul-

tivated fields and wild situations open to the sun, and is adult throughout the

year. Occasionally plagues of this species composed of individuals of a body

size smaller than that of the resident individuals fly over from the African con-

tinent, but on their arrival do not prove noxious.

Acheta meridionalis . : Similar to A. hispanicus, below, but on the average

shorter-winged, this cricket lives in the eastern part of Tenerife, and is es-

pecially common at Barranco Grande. Its habits and seasonal periodism are

similar to those listed for A. hispanicus.

Acheta hispanicus. - This moderate-sized, fully winged cricket lives un-

der stones and debris near the banks of ponds, pools, and dry river beds. It

is essentially southern and eastern in its Tenerifean distribution, and is adult

practically all year.

Modicogryllus guanchicus. : This cricket, smaller-bodied than the preced-

ing and with slightly abbreviated wings, is found along the northern coast of

Tenerife and across the Anaga Peninsula. It is geophilous in humid places, es-

pecially near water. Isolated records of adults are available from each of the

four seasons, so it could well be adult throughout the year.

Modicogryllus palmetorum.. : Another small cricket with slightly abbre-

viated wings, this insect occurs at Barranco Bufadero and Charca de Tahodio,

a southern coastal distribution.

Tartarogryllus burdigalensis. : This widespread Mediterranean species

finds its way to Tenerife, based on one specimen taken at El Medano along the

southern coast.

Platygryllus brunneri. P This species is a moderate-sized, fully winged

geophilous cricket of tropical or subtropical distribution in the Old World. Its

scattered records from Tenerife (Puerto de San Juan to Tahodio) indicate a

southern and western coastal distribution on the island. Its adult seasonal pe-

riodism is uncertain, but could well involve the entire year.

Gryllomorpha canariensis. This small cricket, endemic to Tenerife, is

"ibid. , p

Tenerifean Orthopteroid Distribution 15

apterous. It occurs in a great arc across the interior from near Teno to Ana-

ga, but is never abundant. It appears to be most common in the forests of La

Esperanza and Las Mercedes, where it lives under rocks and debris.

Gryllodes sigillatus. This mottled brown cricket is brachypterous in the

male sex and apterous in the female. It is here recorded from La Esperanza

and Santa Cruz. Several of the Instituto specimens from the latter locality

bear the notations ''En el maiz de la Argentina" or ''Comen maiz Argentina. "'

Mogoplistes squamiger. This is a coastal form found virtually throughout

the Mediterranean area and over to Tenerife and Madeira. It is restricted to

the supratidal, or ''spray"' zone of the sea, where it lives under stones and al-

ae.

Cycloptiloides canariensis. This is a minute, apterous cricket endemic

to Tenerife. It occurs under stones and debris in Santa Cruz and across the

"Laguna Belt" to Barranco Martianez, an essentially coastal distribution.

Trigonidium cicindeloides. This minute, distinctive cricket is widely dis-

tributed in the tropical and subtropical regions of the Old World. In Tenerife

it is phytophilous in cultivated places and in wild, mesic situations within the

"laguna Belt" and over to Puerto de la Cruz on the northern coast.

Oecanthus pellucens, This well-known tree cricket has an essentially

Mediterranean distribution, though it extends into central Europe. It is also

found in Tenerife, where it has a northern and central distribution that, with

additional collecting, could prove to be virtually ubiquitous. It is thamnophi-

lous in habit, and apparently is adult all year long.

Gryllotalpa africana, This is a small species of mole cricket known from

many parts of Africa and Asia, as well as from Spain and the Canaries. It

leads a fossorial existence in humid places along the southern coast of Tene-

rife, chiefly at El] Medano and San Andres.

Paratettix meridionalis. This grouse locust extends throughout the Med-

iterranean region and into the Canaries. In Tenerife it occurs at the banks of

pools and dry river beds throughout the island, and is especially common at El

Medano and San Andres. It appears to be adult throughout the year.

Pyrgomorpha cognata. The presence of this small-bodied pyrgomorph

on Tenerife is based on a record by Gardner (1960), attributed by Holzapfel

(1970) to P. cognata. Inasmuch as Holzapfel lacked access to Gardner's ma-

terial the distributional record supplied (Puerto de la Cruz, along the north-

ern coast) remains uncertain. Nothing else is known with respect to its occur-

rence on Tenerife, but in Spain this geophilous insect frequents arid places o-

pen to the sun.

Arminda brunneri. The genus Arminda, a number of small, apterous

grasshoppers related to the Mediterranean genus Platyphyma (=Pezotettix),

was recently revised by Holzapfel (in press). These interesting insects are

endemic to the Canarian Archipelago, and tend to be restricted to particular

islands. The important species on Tenerife is A. brunnevi. It frequents

16 _ Contrib, Amer. Ent. Inst., vol. 8, no. 1, 1972

shrubby ravines and other elevated land extending from Punta de Tenoto Punta

de Anaga. Here, dwelling principally among laurel forest remnants in excess

of 1,000 meters above sea level it finds its preferred habitat, reaches its great-

est abundance, and is found adult all year long. The insect also extends south-

ward down to the onset of arid coastal scrub, living there on volcanic terrain

with a largely autocthonous vegetation, but never attains great prosperity and

disappears early. Its eclosion seems directly related to rainfall. The first

rains of the season normally. occur in September, and the insect hatches short-

ly thereafter. However, on occasion the rainy season is delayed until as late

as December, in which case it hatches at that time.

Arminda burri. This species is known from Gran Canaria. However, in

the series of Tenerifean Arminda examined by Holzapfel (1970) there was a

male specimen from La Vega that proved indistinguishable from A. burri.

Therefore, the Tenerifean record stands pending further study.

Calliptamus plebeius. This genus was recently reviewed by Jago (1963),

who found only C. plebezus on the Canarian Archipelago. On Tenerife C. ple-

bezus is ubiquitous, inhabiting arid situations ranging in elevation from near

sea level to 2,200 meters above. It is phytophilous on stout herbs and shrubs,

and is encountered in the adult condition largely during the months of June

through October. The insect is striking for its marked sexual dimorphism and

color variants.

Schistocerca gregaria. This large plague locust comes to the Canaries by

invasion from the African continent. The normal winds that affect the island,

the Trade Winds, or Alzszos, are from the northeast, but in October or there-

about they die down, and warm winds from the Sahara Desert, the Harmatan,

may descend upon the islands. In that event, and if there is an outbreak on the

continent, this locust swarms over. During the occasional plagues, as in Jan-

uary, 1954, much of Tenerife is invaded, and thousands of locusts pass over

Santa Cruz like a cloud. At-other times this semidesert species pays only spo-

radic visits. Nevertheless, it is not uncommon to See isolated individuals in

the east during the period October through November.

Scintharista n. notabilis. This xerophilous oedipodine has a southern and

northeastern coastal distribution on Tenerife. It frequents barren lava beds

and other sparsely vegetated situations extending from near sea level to 1, 400

meters above. It is especially common in San Andres, where it spends much

time in flight, producing a strong, clattering crepitation and showing reddish

wings. It is adult throughout the year.

Oedaleus decorus. This is a pale-bodied, yellow-winged grasshopper

strongly banded with dark. On Tenerife it has an interior distribution in dry,

open places from La Laguna to Aguamansa. It is especially common on herbs

in Las Lagunetas. Its adult periodism extends from April through November.

Locusta migratoria. This large plague locust, widely distributed through-

out Africa and extending into Europe and Asia, also occurs in the Canary Is-

lands, where it proves to be in the solitary phase, and is composed of smaller-

bodied individuals than on the mainland. On Tenerife its distribution is a cres-

centic one from Puerto de la Cruz in the north, to Aguamansa, to El Medano

Tenerifean Orthopteroid Distribution 17

in the south, and it appears to be especially common in Los Rodeos and Taco-

ronte. Its habitat consists of cultivated fields, open pasture, and dry shrubby

areas. It isoften highly abundant, but is never noxious. It is in the adult stage

throughout the year.

Oedipoda caerulescens. This grasshopper is listed by Holzapfel (1970) as

occurring in the south and southwest of Tenerife.

Oedipoda canariensis, This xerophile, characterized by pale bluishwing

discs, is endemic to the Canaries. It is essentially a southern and western

coastal form, though it extends considerably inland in the vicinity of the ''La-

guna Belt.'' This species, typical of the genus, inhabits rocky, sandy, or bare

ground exposed tofullsun, but is never very abundant. Records of adult occur-

rence are available for all months except February and March, so it is prob-

ably adult all year long.

Sphingonotus rubescens. 4 Based onthe literature, this species isdescrib-

ed as a darkish, often blotched oedipodine with transparent wings that lack the

typical colored disc outlined by adarkcrescent. It belongs to a genus that, like

Oedipoda, contains many xerophilous geophiles that long have posed a problem.

The group is clearly in need of revision, in connection with which Holzapfel

(1970) indicated the probable necessity of using cytogenetics and behavioral stu-

dies asa supplement to traditional methods of study. She ventured the opinion

that it is presently impossible to separate this species, S. rubescens, from

three others, S. canariensis, S. caerulans, S. savignyi, that have been repor-

ted from the littoral and semi-desert regions of Tenerife at a level generally

not exceeding 300 meters above the sea. Presently we have no recourse but to

take S. rubescens inthe old sense, though excluding it from the central heights

of Las Canadas. Interpreted this way itis virtually ubiquitous, being absent on-

ly from the highest mountains and abounding along the coasts all year long. It

is adult throughout the year, but its maximal presence coincides withsummer.

Sphingonotus savignyi is As presently understood, this contrastingly

marked species has a southern coastal distribution, and it seems to be adult

all year long.

Sphingonotus willemsei 3 According to Holzapfel (1970), thisendemic spe-

cies includes S. teydez. It occursasa geophile in the subalpine desert at ele-

vations of 2,100 meters or above, chiefly during the summer months.

Wernerella pictett. This small oedipodine is a Canarian endemic. On

Tenerife it has an essentially southern and western coastal distribution up to

about 1,000 meters above sea level. Adult records are available from all sea-

sons.

Acrotylus 1. insubricus. This pink-winged grasshopper is widespread

within the Mediterranean region, and extends over to the Canaries. On Ten-

erife it has an expanded coastal distribution in xerophilous situations usually be-

low 650 meters above sea level. It is probably adult throughout the year.

9 ped p. 13.

18 Contrib. Amer. Ent. Inst., vol. 8, no. 1, 1972

Acrotylus longipes. This yellow-winged grasshopper is not listed for

Tenerife by Holzapfel (1970). However, based on Morales Martin's specimens

compared with determined Instituto materials, and others determined by Dirsh,

it is clear that the species is represented in the fauna. It has an essentially

southern coastal distribution, and is adult throughout the year.

Acrotylus patruelis. This small, pink-winged oedipodine is even more

widespread than is A. insubricus, for it occupies roughly the same Mediterra-

nean areas, together with the greater part of Africa. Its distribution within

Tenerife is essentially ubiquitous, though it is most common in dry meadows

in the interior of the island, especially at Aguamansa. It is adult throughout

the year.

Atolopus strepens. This species, a grasshopper widespread through-

out the Mediterranean area, is common throughout the north of Tenerife. It

primarily occupies cultivated fields, pastures, and other open grassy situa-

tions, and is adult throughout the year.

Atolopus thalassinus. Also widespread, both elsewhere and in the Ca-

naries, this species is often collected together with A. strepens. It is espe-

cially abundant at Los Rodeos and Tacoronte, but occurs throughout the island,

except in the central heights. Adults can be taken at any time during the year.

Stenohippus epacromioides. There appears to be but a single species

of Stenohippus on Tenerife. However, the name to use for it is uncertain.

These delicate gomphocerines are variable, difficult to determine, and the lit-

erature confusing. Until more is known it seems best to use S. epacromioides

(the designation used by Uvarov, 1926) in place of S. bonneti (cited by Chopard,

1943, and Holzapfel, 1970). The insect occurs along the southern and western

coasts of the island, and is possibly most abundant near San Andres in the

southeast. It is a graminicole, always being in association with lush grasses.

It is adult all year long.

Doctostaurus maroccanus. This species, a plague locust whose gre-

garious phase has often reached outbreak populations in countries bordering

the Mediterranean Sea, extends well past the Mediterranean world deep into

central Asia. It also occurs, in the solitary phase, in dry, open situations

throughout the north of Tenerife, and is especially common at Las Lagunetas

and San Andres. It probably occurs in the adult stage throughout the year, but,

as indicated by gaps in the record, the adult numbers probably decline mark-

edly during winter.

Truxalis nasuta. This large, distinctive grasshopper has a wide dis-

tribution in the Ethiopian and Mediterranean worlds. It is also known from

Gran Canaria and, based on a record by Gardner, 1960, from Tenerife. If

the latter record is valid, that places it near Puerto de la Cruz. Little else

is known about its Canarian occurrence. However, in view of our knowledge

of its distribution in Spain its habitat is predictable: cultivated and uncultivat-

ed land, often sunny hillsides, where it is always in close association with

grasses for food and perching.

Anataelia canariensis. This small, apterous earwig is the sole spe-

Tenerifean Orthopteroid Distribution 19

cies of a genus of Protodermaptera. The insect is endemic to Tenerife and |

Gomera. On Tenerife it lives along the northern coast from Bajamar to Pun-

ta de Teno, which area constitutes the most ancient terrain of the island.

This distribution, together with the earwig's occurrence on the geologically

similar Playa de San Sebastian of Gomera, suggests that the insect is a relict

form, and opens the possibility that the two islands were united at some time

during the geological past. The earwig's habitat is under stones in the supra-

tidal, or ''spray,'' zone immediately adjacent to the sea, from which area it

never strays more than a few meters (Escalera, 1922, Fernandez, 1966). It

is apparently adult throughout the year, though its numbers become greatly de-

pleted during the summer months.

Labidura riparia. The cosmopolitan Labidura riparia frequents sandy

beaches along the southern coast of Tenerife. It is associated with the under-

side of dry algae, and seems to be adult virtually all year.

Euborellia annulipes. This small, wingless earwig is cosmopolitan

elsewhere in the world, and proves to be widespread throughout the Canarian

Archipelago. On Tenerife it has a crescentic distribution, occurring in the

east of the island and from there across the northern and southern shores. It

lives under stones, often at the edge of the sea, and is adult throughout the

year.

Anisolabis maritima. This earwig, also cosmopolitan and wingless, is

essentially a southern coastal species, though on occasion it proceeds consid-

erable distances inland. Its habits are similar to those of E. annulipes, being

found under stones and often near the sea.

Anisolabis maxima. This large-bodied earwig, like its two relatives,

E. annulipes and A. maritima, is apterous, but it differs from those twocos-

mopolitan species in that it is autocthonous. It occurs in forests throughout

the north of Tenerife, but is most common at Monte de los Silos, Las Canadas

del Teide, and the Anaga Peninsula. It lives under stones, and is adult all

year long.

Labia minor. This minute earwig with abbreviated tegmina is cosmo-

politan in world distribution. On Tenerife it is found along the northern coast

from Puerto de la Cruz to Taganana, with an extension inland at the ''Laguna

Belt.'' It is usually found under stones, debris, rock pile fences, and the foun-

dations of houses.

Perirrhytus edentulus. This small earwig with abbreviated tegmina is

found in Madeira as well as in several of the Canary Islands. On Tenerife it

has a northern distribution, extending most of the way across the island. Adult

records are from winter, spring, and summer, suggesting possible imaginal

absence in fall.

Forficula auricularia. This familiar earwig is Holarctic in distribu -

eo Instituto Espanol de Entomologia record of Anataelia canariensis

from Guimar is probably a mistake, and one from La Laguna is assuredly

incorrect.

20 Contrib. Amer. Ent. Inst., vol. 8, no. 1, 1972

tion. On Tenerife its distributional pattern is crescentic, and with further

study could prove to be circular, encompassing all but the central heights a-

bout Teide. It lives in close association with stones, debris, and other kinds

of shelter, and is adult throughout the year.

Guanchia cabrerae. There are five species in the genus Guanchia, all

from the Canaries, and four occur in Tenerife. They are closely related to

the several brachypterous Forficula that live in the Mediterranean area (Cho-

pard, 1946). Therefore, the genus is interesting more because of its endemi-

city rather than because of any structural specializations. These insects live

exclusively in the woods, especially in laurel forests. They usually inhabit

cracks in the bark of trees and the hollows of trunks, but are also found in dry

leaf litter, especially that of Aeonitum. G. cabrerae is endemic to Gran Ca-

naria and Tenerife, and in the latter occurs in the north, where it seems to be

adult practically all year long.

Guanchia canariensis. This geophilous earwig is endemic to Tenerife.

It has a northern distribution, but extends inland as far as El Portillo. It ap-

pears to be adult during spring and summer.

Guanchia guancharia. This earwig is endemic to Gran Canaria and

Tenerife. In the latter it has aninland distribution in the central and east from

Aguamansa to El Bailadero.

Guanchia uxori. This geophilous earwig, endemic to Tenerife, has a

northern distribution similar to that of G. canariensis. -Adults are recorded

from the winter and spring months. |

Contribution No. 289

Department of Biology

Wayne State University

Detroit, Michigan 48202

U. S. A.

RESUME

Tenerife is the largest island of the Canarian Archipelago, a group of

volcanic islands off the coast of Morocco. It features a mild, uniform climate

and great relief, and supports the growth of diversified biotic communities,

both natural and disturbed. A comparatively rich assemblage of Orthopteroi-

dea finds its home there.

During the last fifty years many reports have been published dealing,

in part, with the Tenerifean orthopteroids, and this literature provides an in-

complete faunal list and scattered notes on habits and distribution. More re-

cently a dissertation appeared (Holzapfel, 1970) that treats the systematics and

distribution of the Canarian Acridoidea, but there is still neither an up-to-date

list of the entire Tenerifean fauna nor a detailed discussion of its biogeography.

The present study, involving analysis of biological data and study of specimens

from several collections, aims toward the partial elimination of that gap in the

literature.

The Tenerifean orthopteroid fauna now numbers 88 species, including

Tenerifean Orthopteroid Distribution 21

13 Blattoidea, 7 Mantoidea, 12 Tettigonoidea, 15 Grylloidea, 29 Acridoidea, and

12 Dermaptera belonging to 31 subfamilies and 58 genera. A total of 24 of the

species (27%)and6 of the genera (10%) are endemic to the islands, and the per-

centage of endemic species per group ranges from as high as 58% (Dermap-

tera) to as low as 13% (Grylloidea), and is positively correlated with aptery or

brachyptery, except in the Acridoidea.

The faunal affinities are mostly North African, but there are also many

general Mediterranean, Ethiopian, and cosmopolitan representatives among

the Tenerifean orthopteroids. A total of 30 of the species are Mediterranean

or Mediterranean-Palearctic in world distribution; 16 species Ethiopian, Med-

iterranean-Ethiopian, or Mediterranean and Afro-Asian; 14 species cosmopol-

itan or tropicopolitan; 4 species Holarctic; and the remaining species endemic.

A total of 15 of the genera prove to be Mediterranean or Mediterranean-Pale-

arctic; 2 genera Ethiopian; 4 genera Ethiopian-Palearctic; 19 genera cosmopol-

itan or tropicopolitan; ll genera globally widespread; and the remaining genera

endemic.

The orthopteroid distributions fall into 37 provisional patterns (explained

in Plates I-IV) that can be aggregated into seven major categories: ubiquitous,

southern, eastern,northern, eastern crescent, interior, and coastal. The

coastal pattern is expanded beyond the usual sense in that it also includes a low

stretch of land, the ''Laguna Belt, '' joining the northern and southern coasts at

a point between the Esperanza and Mercedes mountains. This area is essen-

tially an inland extension of coast, and many coastal species take advantage of

it to live there.

The orthopteroid distributions prove inconsistent with the elaborate bio-

geographic scheme postulated by Ceballos and Ortuno (1951) based primarily on

plant distributions. A better fit of the data is obtained by plotting the distribu-

tions against elevation as modified by exposure. On that basis the following

tentative faunal regions may be recognized: beach, sublittoral scrub, tropical

forest, central forest, and subalpine scrub.

The Tenerifean Orthopteroidea respond to the mild, subtropical Canar-

ian climate in the expected manner. They tend to be active all year long, and

either nymphs or adults may be taken at any time from within the preferred

habitats, except in the subalpine zones. Inthe latter, the inhabitants undergo

a diapause during the winter months, and do not appear as nymphs until spring

and as adults until summer.

RESUMEN

Tenerife es la isla mas extensa del Archipielago de Las Canarias, grupo de

islas volcanicas situado frente a las costas de Marruecos. Sus caracteristicas,

de gran relieve y Suave e uniforme clima,condicionan la formaciOn de diversas

comunidades bioticas naturales e introducidos. Comparativamente es ella

rica en representantes del orden Ortoptera.

Durante los ultimos cincuenta anos se han publicado diversos trabajos que

atafien,solo en parte,a los ortopteros tinerfenos que, en su conjunto, unicamen-

te proporcionan una lista incompleta de dicha fauna y dispersas notas sobre sus

habitats y distribuciones. Recientemente Holzapfel (1, 970) ha publicado un es-

tudio que trata de la sistematica y distribucion de los acrididos de las Canarias,

pero en el no aparece una lista actual de los de la fauna tinerfefia ni una discu-

sion detallada de su biogeografia; el presente estudio, que comprende el anali-

sis de datos biologicos y el estudio de gran numero de ejemplares de diversas

colecciones, aspira, dentro de los posible, a eliminar--aunque solo sea parci-

almente--las lagunas existentes.

22 Contrib. Amer. Ent. Inst., vol. 8, no. 1, 1972

La fauna de Ortopteroides de Tenerife consta hoy dia de 88 especies que in-

cluyen a 13 blatidos, 7 mantidos, 12 tetigonidos, 15 grilidos, 29 acrididos y 12

dermapteros, pertenecientes a 58 generos y 31 subfamilias. Un total de 24 espe-

cies (27%) y de 6 generos (10%) son endemicos de la isla, y el porcentaje de es-

pecies endemicas, por grupos, ofrece maximos del 58% (Dermaptera) y mini-

mos del 13% (Grylloidea), relacionados ellos con el apterismo o braquipterismo

excepto en los Acridoidea.

La afinidad de esta fauna es en su mayor parte con la norteafricana, pero

existen tambien en ella algunos representantes mediterraneos, etiopicos y cos-

mopolitas. Un total de 30 especies son mediterraneas ) mediterrhneas- ‘pale-

arcticas de distribucion mundial; 16 etiopicas, mediterraneas-etidpicas 0 me-

diterraneas y afro- asiaticas; 14 cosmopolitas 0 tropicales; 4 holarcticas y el res-

to especies endemicas. 15 de sus generos son mediterraneos 0 mediterraneos-

palearcticos; 2 etidpicos; 4 etidpicos- -palearcticos; 19 cosmopolitas 6 tropicales;

ll de dispersion mundial y el resto endemicos.

Con estos taxones se han hecho 37 grupos de tipo provisional (expuestos en

las laminas I a IV) y reunidos, en las siete siguientes categorias principales:

ubicuos, surenos, orientales, nortenos, orientales ascendentes, interiores y

costeros. El grupo costero comprende algo mas de lo que indica la acepcion

usual de ese termino ya, que incluye el una baja y larga faja de tierra (Paso de

La Laguna), que une las costas norte y sur por un paso situado entre los mon-

tes de La Esperanza y Las Mercedes; esta area por ser esencialmente una ex-

tension de la costa, es aprovechada por varias especies costeras para vivir en

ella.

La distribucion de estos Ortopteroides muestra la poca consistencia que pa-

ra ellos ofrece el esquema biogeografico aportado por Ceballos y Ortuno (1, 951)

basado, principalmente, en la distribucion de las plantas. Trazando su distri-

bucion ‘sobre un mapa que descanse sobre las zonas altitudinales afectadas por

la orientacion, se consigue un mejor aprovechamiento de los datos existentes

y con base en ello, intentamos poner de manifiesto las siguientes zonas faunis-

ticas: litoral, sublitoral, bosque tropical, bosque central y zona subalpina.

Los Ortopteroides de Tenerife responden, como era de esperar, a las ca-

racteristicas del suave y subtropical climadel archipielago canario. Suelen ser

ellos activos durante todo el ano, y tanto ninfas como adultos pueden colectarse

en cualquier epoca en sus habitats preferidos, excepto los de la zona subalpina.

En esta ultima, sufren una pausa durante los meses invernales y no aparecen

sus ninfas hasta la primavera y en verano los adultos.

ACKNOWLEDGMENTS

The data reported herein were amassed during the tenure of the first au-

thor's grants from the American Philosophical Society and the Graduate Divi-

sionof Wayne State University. Insect specimens were loaned by Sr. Dn. R. Agen-

jo, Director, and Srta. Dna. Vicenta Llorente del Moral, Orthopterist, of the

Instituto Espanol de Entomologia, Madrid, and by Sr. Dn. Jose Maria Fernan-

dez, Chief, Laboratorio de Zoologia, Museo de Ciencias Naturales, SantaCruz.

Sres. Fernandez, above, and E. R. S. Sventenius, Jardin de Aclimatacion de

la Orotava, Puerto de la Cruz, assisted in certain plant determinations. Sres.

Manuel Garcia de Viedma e Hitos, Catedratico, Escuela Tecnica Superior de

Ingenieros de Montes, Madrid, Jose Miguel Gonzalez Hernandez, Chief, Pat-

Tenerifean Orthopteroid Distribution 23

rimonio Forestal del Estado, Santa Cruz, and Salvador Vazquez Duque, Las

Mimosas, Santa Cruz, gave advice and rendered various other professional

services essential to the project. Miss Leslie A. Gangwere assisted in col-

lecting specimens in the field. Irving J. Cantrall, Curator, Insect Division,

University of Michigan, Ann Arbor, and William L. Thompson, Professor,

and William Prychodko, Associate Professor, both of the Department of Biol-

ogy, Wayne State University, Detroit, read the manuscript and offered sugges-

tions of value in its improvement. Mrs. Betty Kuehn, also of the Department

of Biology, Wayne State University, typed the manuscript. To the above indi-

viduals and organizations the authors are indebted.

REFERENCES

Ceballos, L., and F. Ortuno.

1951. Estudio sobre la vegetacion y flora forestal de las Canarias Oc-

cidentales. Inst. For. Invest. Exper., Madrid. 465 pp.

Chopard, L.

1942. Insectes Orthopteroides recoltes dans les iles Atlantiques.

Comm. Biol. Soc. Sci. Fenn., 8: 13 pp.

1943. Orthopteroides de l'Afrique du Nord. Librairie Larose, Paris.

450 pp.

1946. Les Orthopteroides des iles Atlantides. Mem. Soc. Biogr.,

8: 199-208.

1954, Insectes Orthopteroides recoltes aux iles Canaries par M. H.

| Lindberg. Comm. Biol. Soc. Sci. Fenn., 14: 15 pp.

Escalera, F. M.

1922. Nota biologica sobre la Anataelia canariensis1. Bol. de Tener-

ife. Bol. R. Soc. Espan. Hist. Nat., 22: 157-158.

Fernandez, J. M.

1966. La fauna entomologica Canaria y sus origenes. Publ. Mus. Ar-

queol. Cabildo Ins. Tenerife, 6: 61-68.

Gangwere, S. K.

In press. Feeding behavior in the orthopteroid insects of Tenerife, Ca-

nary Islands, Spain. Yearbook Amer. Phil. Soc. 1970.

Gardner, A. E.

1960. Odonata, Saltatoria and Dictyoptera collected by Mr. E. S. A.

Baynes in the Canary Islands, 1957 to 1959. Entomologist,

93: 128-131.

Holzapfel, C. M.

1970. Zoogeography of the Acridoidea in the Canary Islands. Univ.

Mich. doct. diss.: 213 pp.

In press. Evolution in the Canary Islands. III: Two new Arminda,

with notes on the Arminda brunneri complex on Tenerife. Occ.

Papers Mus. Zool. Univ. Mich.

Jago, N. D.

1963. A revision of the genus Calliptamus Serville. Bull. Brit. Mus.

Nat. Hist., 13: 289-350.

Johnsen, P.

1970. Notes on African Acridoidea in Danish museums. Nat. Jutland.,

15: 123-162.

24 Contrib. Amer. Ent. Inst., vol. 8, no. 1, 1972

Lems, K.

1960. Floristic botany of the Canary Islands. Sarracenia, 5: 94 pp.

Uvarov, B. P. |

1926. Grasshoppers from Northern Nigeria. Trans. Ent. Soc. London,

1925: 413-453.

1948. Tettigoniidae and Acrididae collected in 1931 on the Atlantic Is -

lands by R. Frey and R. Stora. Comm. Biol. Soc. Sci. Fenn.,

8: 7 pp.

Willemse, C. :

1936. Une excursion orthopterologique auxilesCanaries. Nat. Maandbl.,

25: 40-42, 56-57, 72-73, 86-89, 101-103, 113-115.

1949, Addition to the knowledge of the Dermaptera and Orthoptera of the

Canary Islands. Tijdschr. voor Ent., 91: 7-ll.

1950. Second addition to the knowledge of the Dermaptera and Orthop-

tera of the Canary Islands. Tijdschr. voor Ent., 92: 248-250.

Willemse, C., andC. F. A. Bruijning.

1949. On a small collection of Orthoptera from the Canary Islands.

Tijdschr. voor Ent., 91: 23-24.

Tenerifean Orthopteroid Distribution

TABLE I

Annotated Provisional List of Orthopteroidea of Tenerife, Canary Islands.

World Distribution

& Endemism

Groups & Species

BLATTOIDEA

Ectobiinae

Arbiblatia bivittata (Brulle) Yes:

Tenerife & G. Cana

ia

Pseudomopinae

Blattella germanica (L.) No:

Cosmopolitan

Loboptera canariensis (Chop. 9 Yes:

Tenerife

Loboptera decipiens (Germ. ) No:

Mediterranean

Loboptera fortunata Kr. Yes:

Tenerife & La Palma

Blattinae

Blatta orientalis L.° No:

Cosmopolitan

Periplaneta americana (L. ) No:

Cosmopolitan

Periplaneta australasiae (Fab. )5 | No:

Cosmopolitan

Periplaneta brunnea Burm.® No:

Cosmopolitan

Oxyhaloinae

Leucophaea maderae (Fab. ) No:

Cosmopolitan

Pycnoscelidinae

Pycnoscelis surinamensis (L. ) No:

Cosmopolitan

Tiviinae

Zetha vestita (Brulle) No:

Neotropical, Maca-

ronesian, Hawaiian

Tenerifean

Distribution?

125.6, 9.8, 16

26, 29, 43, 49,

53, 77, 78, 79,83,

92, lll, 114, 116,

7, 128

60, 37, 102, 116,

124

96, 102

3, 29, 60, 77,

14

7, 18, 49, 60, 116

60, 73

60, 73, 83, 87,

102

60, 102, 120

60

60, 73, 87, 102

14, 32, 43, 66,73,

83, 87, 202, 102,

104, 114, 117, 122

14, 60, 98, 106

Tenerifean

Distribution

Type®

IVa

Vb

vilt

Vie

IVd

Vili

VIlid

Vilt

Vilf

Vild

Vila

Vile

29

Adult

Seasonal

Periodism

All months ex-

cept VII & XII

I-IV, VII-VIIL,

XX

IV, VI-VII

All months ex-

cept I, VI, VII,

XII

x, wit

I, IV-V, VII-XI

i; Xx

VI

i Vil, 1X, x1

I-III, V-VI

All months ex-

cept IV, X

26 Contrib. Amer. Ent. Inst., vol. 8, no. 1, 1972

TABLE I, CONT.

| Tenerifean Adult

World Distribution Tenerifean Distribution Seasonal

Groups & Species & Endemism® Distribution? Type® Periodism9

Epilamprinae

Leurolestes ciycumvagans Burm. | No: B85, 102 Vili I, V, XI

Cosmopolitan

MANTOIDEA

Amelinae

Ameles gracilis (Brulle) No: D,) 2d,aOn Od VIb VI-VII, IX-X_

Mediterranean .

Ameles limbata Brulle No: (12; TA 204/36) Ic IV-VI, IX-XI

Mediterranean D1, 64, 73, 93,117,

128

Pseudoyersinia subaptera Chop. Yes: 22, 35, 43, 46,66, VIith I, II-VI, XI

(=P. lindbergi Chop. ) Tenerife & Lanzarote {79, 81, 117

Pseudoyersinia teydeana Chop. Yes: 18 Vig III, VI, VIII

Tenerife

Mantinae

Mantis religiosa (L.) No: 3, 5, 60, 89, 91, Ve VI-XI

Palearctic & Nearctic|1l4, 124

Empusinae

Blepharopsis mendica (Fab. ) No: 55343505, 13,88, VIIb III-IV, VI, VIII-X

Mediterranean 102, 122, 124

Hypsicorypha gracilis (Burm. ) No: 7, 14, 48, 60, 66, Vb II-VIII

N. African 73, 83, 100, 102,

103, lll, 123

TETTIGONOIDEA

Phaneropterinae

Phaneroptera nana sparsa Stal No: do, 9, 14, 18, 22, Ia All months ex-

Mediterranean & 43, 49, 51, 55,58, cept X

Ethiopian 09, 60, 87, 88,

100; 102,277, 7

Meconeminae

Canariola nubigena (Kr. ) Yes: Dy t,t, OT GT, IIIb II-IV, VI

Tenerife 7

Canariola willemsei Mor. Yes: 5, ll4 | Vid IX, XI

Tenerife & Gomera

Tettigoniinae

Tenerifean Orthopteroid Distribution 27

TABLE I, CONT.

Tenerifean Adult

World Distribution Tenerifean Distribution Seasonal

Groups & Species & Endemism Distribution Type Periodism

Calliphona k&nigi Kr. Yes: By By Dy ey eG; IVec IV-IX, XII

Tenerife 39, 46, 60, 72,

83, 94, lll, 116,

119

Decticinae

Ariagona margaritae Kr. Yes: Le a 7, 13, IVe VII-XI

Tenerife, Hierro, & | 29, 57, 63, 65,

Gomera Tt, 209, Tb 415

? Platycleis denticulata Panz. No: ---- ---- ----

W. Mediterranean

Platycleis falx (Fab. ) No: ---- ---- ----

Mediterranean

Platycleis grisea (Fab. ) No: 26, 114 Vif VI, IX

Palearctic

Platycleis intermedia (Serv. ) No: 14, 23, 67 VIIf IV-V, VII

Palearctic (especially

Mediterranean)

Platycleis sabulosa Azam No: o,. 7, 14; 23, 25, IVc II-VI, [X-X

W. Mediterranean 49, 60, 69, 77,

Bo, 87, M0> ¢24;

117

Tessellana tessellata (Charp. ) No: BG 80. Ore 77. Illa IlI-V, VII-IX, XI

Mediterranean & Cal-| 83, 91, 96, 724,

ifornian 117

Decticus albifrons (Fab.) No: Duty 20500, 266, Illa IlI-IX, XI

Mediterranean 87, 110, “U2, 17

GRY LLOIDEA

Gryllinae

Gryllus bimaculatus De G. No: 5, 8, 14, 26, 28, IVb All months

Afro-Asian & Medi- 43, 51, 60, 64,

terranean 6G, 69, 70" 77,

83.87, 89, LOL,

102, 111, 114, 116,

125

Acheta meridionalis (Uv. ) No: 14, 24, 27, 37, IIIc All months ex-

(=A. canariensis Chop.) Mediterranean 43, 60, 94, 102, cept VI-VIII

114, 7, 122

Acheta hispanicus (Ramb. ) No: 5, 7, 14, 43, 60, IIc IlI-V, VII-XII

Mediterranean 73, 114

28 Contrib. Amer. Ent. Inst., vol. 8, no. 1, 1972

TABLE I, CONT.

Tenerifean Adult

World Distribution Tenerifean Distribution Seasonal

Groups & Species & Endemism 6 Distribution? Type8 Periodism9

pe a ie doa burdigalensis No: 73 Vij VII

(Latr. ) Mediterranean

Modicogryllus guanchicus (Kr. ) No: 7, 14, 24, 60, 90, IVd II, IV-VI, VIII,

Spanish Sahara 114, 117, 118 ».4

Modicogryllus palmetorum (Kr.) | No: 17, 118 VI II, IV-V, IX

Mediterranean &

Ethiopian

Platygryllus brunneri (Sauss. ) No: Ba Ot, 0k, Ti; VIIh I, I-IV, VII, IX

Almost tropicopolitan}118

Gryllomorpha canariensis Chop. | Yes: 3,. 18, 26, 27, (28, via I, VI, IX-XI

Tenerife 45, 52, 64, 67,

lll, 116, 117

Gryllomorpha longicauda {(Ramb. ) | No: ---- ---- ----

Mediterranean (Iber-

ian & N. African)

Gryllodes sigillatus (W.) . No: 26, 102 IlId VI, XI-XII

Tropicopolitan

Mogoplistinae

Mogoplistes squamiger (Fisch.) | No: 102 VII1 I-II, IX

Mediterranean &

Madeiran

Cycloptiloides canariensis (Bol..) | Yes: 60, 69, 102 Vile I, XII

Tenerife

Trigonidiinae

Trigonidium cicindeloides (Ramb.)| No: 60, 87, 109 Vile H,- Vi, &

Afro-Asian & Medi-

terranean

Oecanthinae

Oecanthus pellucens (Scop. ) No: Ds. Once alan GO, IVe II, V, VII-XI

Palearctic, especial-|64, 83, 87, 205,

ly Mediterranean 124, 128

Gryllotalpinae

Gryllotalpa africana (Beauv. ) No: 5a,.°735:-93 VIli IV, VI, X

Mediterranean &

Ethiopian

Groups & Species

ACRIDOIDEA

Tetriginae

Tetrix subulata (L. )

Paratettix meridionalis (Ramb. )

Pyrgomorphinae

? Pyrgomorpha cognata Kr.

Catantopinae

Arminda brunneri Kr.(=A. appen-

hageni End.)(= A. striatifrons

End. )

? Arminda burri Uv.

Calliptaminae

Calliptamus plebeius (Walk. )

Cyrtacanthacridinae

Schistocerca gregaria (Forsk.’)

Oedipodinae

TABLE I, CONT.

World Distribution

& Endemism®

No:

Holarctic

No:

Mediterranean

No:

Mediterranean &

Ethiopian

Yes:

Tenerife

Yes:

Tenerife & G. Cana-

ria

Yes:

Tenerife, G. Cana-

ria, La Palma, Hier-

ro, & Gomera

No:

Afro-Asian & Medi-

terranean

Tenerifean

Distribution?

Tenerifean Orthopteroid Distribution

5, 43, 47, 53, Ib

60,

114,

128

69

2, 5, 7, 26, 27,

38,

77,

96,

116,

127

73, 87, LOL,

117, 118, 224,

Vilg

IVa

46, 49, 64,

85, 87, 93,

lll, 114, 115,

117, 126, 128

5, 7, 14, 15, 26, la

18, 19, 22, 23,

30, 40, 43,

46, 49, 53,

60, 62, 65,

73, 80, 82,

85, 86, 87, 88,

93, 94, 110,

115, 117, 124,

4, 31, 60, 69, Ve

73,

83, 102

Tenerifean

Distribution

Type 8

29

Adult

Seasonal

Periodism9

I, II-V, VIII,

X-XII

II

All months

I, VI-VIII, X

I-Il, IX-XI

30 Contrib. Amer. Ent. Inst., vol. 8, no. 1, 1972

TABLE I, CONT.

Tenerifean Adult

World Distribution Tenerifean Distribution Seasonal

Groups & Species & Endemism® Distribution Type Periodism

Scintharista n. notabilis (Walk.) | No: 9, 19, 42, 43, IIb All months

Mediterranean & 53, 54; 55, 73,

S. Asian 93, 94, 102, 110,

Lt, 128

Oedaleus decorus (Germ. ) No: 3, 45°°5,--29,- 60, Vid IV, VI-VIII, X-XI

Mediterranean & 65, 91

Ethiopian

Locusta migratoria (L. ) No: 3. 53) 14. 19. 23, Ve All months

Afro-Asian & Medi- 28, 37, 39, 43,

terranean 46,60, 69, 73,

Ui 19s 87s SS,

91, 93, 94, 102,

105, W445 115; 157

Oedipoda caerulescens (L. ) No: 2, 56,928 IIb ----

Mediterranean

Oedipoda canariensis Kr. Yes: 1, 19, 22, 46, 60, IIb All months ex-

Tenerife, G. Canar- | 64, 73, 79, 82, cept II-III, VII

ia, La Palma, Go- 03,-°94,°202, 117

mera,& Azores

? Oedipoda fuscocincta Luc. No: ---- ---- ----

N. African

Sphingonotus caerulans (L. ) No: 71 ---- ----

Palearctic

Sphingonotus canariensis (Sauss.) |No: 22 VIIk IV

Mediterranean (N.

African)

? Sphingonotus lucasii Sauss. No: ---- ---- ----

Mediterranean (N.

African)

Sphingonotus rubescens (Walk.) _|No: R, lds 1: 28597. Ib All months ex-

Afro-Asian, Mediter-| 32, 35, 43, 51,54, cept VII

ranean, & Madeiran 6G,09,- Ta, (9,

- 7-87, 88, 94, 102,

108, 110, lll, 117,

128

Sphingonotus savignyi Sauss. —_ |No: 31, 44, 78, 74, Vili I, II-IV, VI-IX,

Mediterranean 117 XI

Sphingonotus willemsei Mists. Yes: 18 vig VIT-IX

(=S. teydez) Tenerife

? Wernerella aspera (Brulle) No: ---- ---- <=

Mediterranean

Groups & Species

Wernerella picteti (Kr.)

Acrotylus i. insubricus (Scop. )

Acrotylus longipes Charp.

Acrotylus patruelis (H.-S. )

Aiolopus strepens (Latr. )

Atolopus thalassinus (Fab. )

Gomphocerinae

Stenohippus epacromioides

Krauss (=S. bonnet)

Dociostaurus maroccanus (Thunb.

Truxalinae

Tenerifean Orthopteroid Distribution

TABLE I, CONT.

World Distribution

& Endemism

Yes:

Tenerife & Gomera

No:

Mediterranean

No:

Mediterranean

No:

Mediterranean &

Ethiopian

No:

Mediterranean &

Ethiopian

No:

Mediterranean &

Ethiopian

No:

Ethiopian & Cape

Verdean

No:

Mediterranean

Tenerifean

Tenerifean Distribution

Distribution? Type’

19, 32) 2a 27 IIb

43, 46, 55, 70,

73

th, Woe BU Res, Vila

54, 55,66, 69,

87, 88, 98, 102,

LHL Sap ies

43, 73, 75, 128 IIc

S, 7,-44, 519, 23, Ib

26, 28, 29, 37,

43, 51, 60, 64,

69, 73, 77, 83,

87, 88, 93, 94,

99, 100, 102, 205,

107, 110, 114, 117,

128

2: 3. Be 7 ai AS, IVa

16, 18, 28, 27,

28, 43, 46, 51,

55, 60, 64, 69,

U1, 88, 87, 88,

93, 94, 99, 202,

102, 205, 111, 114,

116, 117, 228

%, 22, 23, 29.32, Ib

37, 43, 49,51,

55, 60, 69, 77,

83, 87, 88, 90,

91, 93, 94, 99,

102, 109, 110,222,

114, 116, 117, 120,

124

19, 34, 66, 73, VIIh

76, 98, 94, 95,

102, 117

5, 23, 26, 29, IVb

46, 53, 60, 65,

70, 77, 87, 88,

93, 110, 114, 117

31

Adult

Seasonal

Periodism

I, II-VI, VII-x,

XII

I, I-IV, VI-VIIL,

XI-XII

[) TSE; VI;

VIll-X, XII

All months

All months

All months

Ali months ex-

cept II, VII

IlI-IX, XII

32

TABLE I, CONT.

Contrib. Amer. Ent. Inst., vol. 8, no. 1, 1972

Groups & Species

? Truxalis nasuta (L.)

PROTODERMAPTERA

Anataelia canariensis Bol.

DERMAPTERA

Labiduridae

Labidura riparia (Pall. )

Euborellia annulipes (Luc. )

Anisolabis maritima (Gene)

Anisolabis maxima (Brulle)

Labiidae

Labia minor (L. )

Forficulidae

Perirrhytus edentulus (Woll. )

Forficula auricularia L.

World Distribution

& Endemism

No:

Mediterranean &

Ethiopian

Yes:

Tenerife & Gomera

No:

Cosmopolitan

No:

Cosmopolitan

No:

Cosmopolitan

Yes:

Tenerife, G. Canar-

ia, Hierro, & Gomera

No:

Cosmopolitan

Yes:

Tenerife, G. Canar-

ia, Gomera, & Ma-

deira

No:

Holarctic

Tenerifean

Distribution’

69

14, 41, 87, 125

31, 73, 84

3, 5.9, a4.

23, 26, 29, 50,

53, 60, 64, 69,

73, 77, 83, 84,

87, 101, 102,109,

114, 16, 117

bi $8y 203,

73, 77, 84, 93,

102, 113, 116

s. 8. 7, 8

14, 15, 18, 26,

28, 60, 65, 77,

102, 207, 109,

ll, 114, 15, 16,

17

14, 60, 87, 88,

115

5, 33, 60, 115

a5 Aa,

26, 28, 48, 49,

50, 60, 73, 77,

102, 105,207, 122

113, 114

Tenerifean

Distribution

Type

Vilg

Vilg

VIlj

IVa

Vile

Adult

Seasonal

Periodism

II

All months

I, II-VI, VIII-IX,

XII

All months ex-

cept VII

Wi<V, DEX, Ri

All months

| II, VI, XI-XII

I, II-VI, VIII

All months ex-

cept VII-VII

Groups & Species

Guanchia cabrerae (Bol. )

Guanchia canariensis Burr

Guanchia guancharia (Hell. )®

Guanchia uxori (Hell. )

5

Species whose distribution, as shown here, is open to question, being based on 10 or fewer specimens.

6

If yes, the species is endemic, and the Canarian islands where it occurs are indicated; if no, it is not

Tenerifean Orthopteroid Distribution

TABLE I, CONT.

World Distribution

& Endemism

Yes:

Tenerife & G. Canar-

ia

Yes:

Tenerife

Yes:

Tenerife & G. Canar-

ia

Yes:

Tenerife

Tenerifean

Tenerifean Distribution

Distribution” Type®

2,7, 13, 14, 20, IVd

28, 33, lll, 115,

116

2, 7, 77, 85, 92, Via

ll, 15, 16

5, 18, 26, 77, Vic

117

oS & € 12, Via

26, 65, 68, 85,

92, lll, 15

a Canarian or Macaronesian endemic, and its world distribution is noted.

7

The distributional records here given are based on the collections of M. Morales Martin, those of the

Laboratorio de Zoologia of the Museo Insular de Ciencias Naturales, Santa Cruz, and those of the Instituto Es-

pafiolde Entomologia, Madrid, except for the italicized records, which are from literature cited. The numer-

ical code to distribution appears on pp. 4-5.

8

33

Adult

Seasonal

Periodism

All months ex-

cept VII-VIII,

XI-XII

III-VII

II, V-VII, X

I-VI

The explanation of distributional types here used is given on pp. 7-8 and in Plates I-IV.

°The months in question are given in Roman numerals from I (January) through XII (December).

Contrib. Amer. Ent. Inst., vol. 8, no. 1, 1972

34

1seO0

9¢ 968 “Z 9 “PG © uit 6 “GCE 97 ULBSB9Y NOG ZNIf) BUS

1g ple ‘€ G°LT L*§ 8 6r9 LOS °2 [erzUID BueZ]

(..J19@ Bunse’T,,)

eo DLE G c 6L 8 TI O°LLL 1v9 [etyu9o SCY sospoy so'y

(.W19@ eunse’yT,,)

v9 912 ‘Z I “TZ €°CI vV “89S LYG Teruo. {Sey eunseyT eT

IIT SoS 6 “GC VOT 0907 OOT }SBO9 ULOYILON BABIOIO BT

rea xX /sheq ung Jo | (°Do UT) (*Do UT) ( °ULUL UT) (° Ur UT)

JSVIIOAQ “SUH [enuuy ‘“dwiady, "dura, [[eyurey UOTICVASTY UOTS9Y AVID

“AV TRIOL “AVIeIOL ‘“Sny ‘Ay ‘qeq ‘Ay yenuuy ‘ay

“(IG61) OUNZIOQ pue SOTTeQaD pue ‘(qggT) BULdSY] Op [eUOTOEN S¥TIV

‘(8961 ‘9 “IOA) Sp1od9y AaYyyVOM P[LIOM J9IFV ‘SOTIITVIOT UvasIIOUaT, poyoEeTEeg Jo Vyeq [BVITSOTO109}0/

Il Hla VL

Tenerifean Orthopteroid Distribution 5)

SEP OC<HIOCNSHSO

Se enan

“8

~

>4e

WA

ole |

— Mee let of

autine a, UE P2OS

Ne EE

BPX ISe. _~«m

wr, Gel OCR CHS SMD

Hefee

PLATE I. Orthopteroid distributions in Tenerife, as recognized in this re-

port: Distribution Types I-III. ?

Contrib. Amer. Ent. Inst., vol. 8, no. 1, 1972

36

Orthopteroid distributions in Tenerife, as recognized in this re-

port: Distribution Types IV-V.

PLATE II.

Tenerifean Orthopteroid Distribution 37

| i

aca Sorte

PA

OTT ee ie

sRUBRRTERO REMY,

Oe

PLATE III. Orthopteroid distributions in Tenerife, as recognized in this re-

port: Distribution Types VI-VII.

Contrib. Amer. Ent. Inst., vol. 8, no. 1, 1972

38

VII F

PLATE IV. Orthopteroid distributions in Tenerife, as recognized in this re-

port: Distribution Types VII, Continued.

Tenerifean Orthopteroid Distribution 39

TACORONTE

LA OROTAVA

PUERTO DE LA CRUZ

LA LAGUNA

“CANDELARIA

GUIMAR

IZANA

PUERTO DE SAN JUAN

LOS CRISTIANOS VILAF LOR

COASTAL SCRUB

ites > Bes) SABINAR

LAURISILVA & FAYAL-BREZAL

PINAR

ESCOBONAL

RETAMA - CODESO

VIOLETA

PLATE V. A. Elevational zones and selected localities of Tenerife. The

encircled dots indicate places listed in Table II. B. Phytogeo-

graphic regions of Tenerife, after Ceballos and Ortuno (1951).

40

Contrib. Amer. Ent. Inst., vol. 8, no. 1, 1972

PUNTA HIDALGO

AFUR

TAGANANA

TACORONTE | BAJAMAR \ TA LAGUNA ay,

a a ,

PUERTO DE LA CRUZ TEJINA r 3 8

7

SAUZAL . 20 77

. 54-—-”—~ IcuESTE

SANTA URSULA 78/ 57 IL

39 . SAN A

SAN JUAN DE LA RAMBLA “tg 91 |? a 58 79 (ie ae sci

A #99 39 30 e SANTA CRUZ

REALEJO a ee LA CUESTA

BUENAVISTA Si ik ata cas a 106 id fe

Boros 64-63

TENO y | BARRANCO GRANDE

‘ 89 e ° 59

_)—_77] 72 SAN DIEGO

« ov s

2 ” 33 497 . LA ESPERANZA

15 45 46 BARRANCO Hi

70 at 40 fe eee

3d 36 CANDELARIA

52 25

107 jee ARAFO

LOS GIGANTES ———/” 18 GUIMAR

e

‘ 80

38 AGUAMANSA

CHIO

-%

PUERTO DE SAN JUAN

PLATE VI.

36 ARICO VIEJO

55

e Ag

—\

‘i ee

LOS CRISTIANOS EL MEDANO 4

wie

|

PORIS DE ABONA

ADEJE

LAS GALLETAS

LA CUESTA

A. Collection localities of Tenerife, as cited in this report. The

location of settlements is indicated by dots, and the name of each

is written out, while the approximate location of other places is

indicated by number. For a code to all locality numbers see

pp. 4-5. The insert below (B) represents a magnified portion

of coast in the Santa Cruz area.

ht

FA 3

ee

f 5 yi

inte

i 4

+

+

*

Dee RT

As a eS

Contributions

of the

American Entomological Institute

Volume 8, Number 2, 1972

MOSQUITO STUDIES (Diptera, Culicidae)

XXVII. The varipalpus group of Aedes (Ochlerotatus)

By J. Hal Arnell and Lewis T. Nielsen

MOSQUITO STUDIES (Diptera, Culicidae)

XXVIl

THE VARIPALPUS GROUP OF AEDES (OCHLEROTATUS)*

By

J. Hal Arnell? and Lewis T. Nielsen®

The Varipalpus Group of Aedes consists of 5 species of treehole-breeding mos-

quitoes with a distribution through much of western North America west of the

Continental Divide. Coquillett (1902:292-293) described Culex varipalpus from a

single female collected at Williams, Arizona. Ludlow (1905:231-232) described

Taeniorhynchus sierrensis from material sent from Three Rivers, Tulare Co., Cal-

ifornia, and subsequently (1906:132) reduced it to synonomy with varipalpus.

The ‘“‘western treehole mosquito” of Arizona and the Pacific Coast was consid-

ered to be Aedes varipalpus until 1956 when Belkin and McDonald (1956:344)

resurrected sierrensis for the populations from the Pacific Coast States and sub-

sequently described (1957:179-191) monticola from southern Arizona. A fourth

species in the Varipalpus Group, deserticola, was described by Zavortink (1970: 1-

7) from the desert of southern California.

Despite the widespread distribution of the Varipalpus Group and the consid-

erable economic importance of sierrensis, a major pest species in localized areas

of California, the group has never been the object of a thorough taxonomic study.

The biology of sierrensis from California has received considerable attention but

the distribution and systematics of the group are poorly known.

We are indebted to John N. Belkin for guidance and encouragement during the

study and preparation of the manuscript; to Thomas J. Zavortink for invaluable

assistance in securing specimens from southern California and Arizona, the pre-

liminary work on deserticola and for advice and suggestions freely given during

many hours of discussion of the study; to Jay H. Linam, Robert J. Novak and

Steven Romney for assistance and companionship in the collecting of specimens;

to William A. Powder and Sandra J. Heinemann for assistance in preparing mater-

ial for study; to Margaret Kowalczyk and Nobuko Kitamura for preparation of

preliminary drawings and inking the final plates; and to Caryle Stallard for the

typing of preliminary drafts, editorial assistance and preparation of the text copy

‘Contribution 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 by J. Hal Arnell

to the University of Utah and revised by him during tenure of postdoctoral traineeship at the

University of California, Los Angeles (U.S. Public Health Service Training Grant 5-T01-AI-00070).

* Department of Zoology, University of California, Los Angeles, California 90024. |

* Department of Biology, University of Utah, Salt Lake City, Utah 84112.

2 Contrib. Amer. Ent. Inst., vol. 8, no. 2, 1972

for lithoprinting. We are also indebted to the U.S. National Museum and the British

Museum (Natural History) for the loan of specimens examined during this study:

MATERIAL AND METHODS

MATERIAL. The material for this study came primarily from collections made

by the authors from April 1965 to October 1970 and material collected for the

project “Mosquitoes of Middle America” (Belkin, Schick, et al., 1965) deposited

at the University of California at Los Angeles [UCLA]. Material from the U.S.

National Museum [USNM] and specimens of the Aedes pulchritarsis group from

the British Museum (Natural History) were also examined. The material collected

by the authors will be deposited at the University of Utah [UTAH] and UCLA.

Several collectors are identified in the distribution lists by initials only. These are

Thomas J. Zavortink (TJZ), Robert J. Novak (RN), Dennis W. Heinemann (DWH),

J.H. Linam (JHL), L.T. Nielsen (LTN) and J.H. Arnell (HA). A total of 15,776

specimens of all species was examined, 2,687 males, 2,548 females, 3,215 pupae

and 7,326 larvae. Of this material, 1,172 were individual rearings (946 larval, 182

pupal, 44 incomplete). Material was collected and prepared for study primarily

following the techniques outlined by Belkin (1962) and Breland (1957).

TAXONOMIC PROCEDURE. Methods used in this study were primarily those

of classical comparative morphology. The material was examined in detail in all

stages and compared with topotypic material of previously described species to

determine its taxonomic position. Group and species descriptions in general fol-

low the method of presentation of Belkin (1962). Terminology for the descrip-

tion of the mesonotal scale patterns is after Berlin (1969). Usually at least 10

adults of each sex from each collection were examined in complete detail with

the remainder examined in pertinent details only. The complete chaetotaxy of

5 to 10 larval and pupal skins from each collection was examined when material

was available and the remainder examined in pertinent details only. All stages of

each species are illustrated. The larval and pupal chaetotaxy figured represents

the modal condition of at least 10 topotypic specimens, as do the illustrations

of the male genitalia. Details of the adults of varipalpus are illustrated as repre-

sentative of the entire group,. with only the mesonota of the remaining species

illustrated.

SYSTEMATICS

TAXONOMY. The Varipalpus Group can be distinguished from all other New

World Aedes as adults by the white scale patches on the palpus and apical white

rings on the tarsal segments. The larvae are characterized by a poorly-developed

ventral brush composed of 6 pairs of hairs with each hair usually double and be-

coming stronger distally.

The 5 species of the Varipalpus Group can be divided into 2 subgroups on

the basis of morphology, 1 represented by laguna, monticola, varipalpus and des-

erticola, the other by sierrensis, the differences between the 2 subgroups being

most evident in the male and larva. In the male, laguna, monticola, varipalpus

and deserticola are characterized by a ventral white streak on the basal 0.5 of

the proboscis, which sierrensis lacks, and the genitalia are simple with relatively

Arnell & Nielsen: Aedes varipalpus Group 3

minor variations among the species in the basal tergomesal lobe of the sidepiece

and the lobes of the ninth tergite. The male genitalia of sierrensis have a rela-

tively short, broad sidepiece, the basal tergomesal lobe with a dense cluster

of strongly developed setae rather than a single seta, short strong hairs on the

tergomesal margin of the sidepiece, a relatively short clasper, and a relatively short

claspette. In the larva, laguna, monticola, varipalpus and deserticola differ from

each other primarily in rather slight variations in chaetotaxy and siphon length,

while sierrensis differs from these in gray instead of white coloration, weaker ab-

dominal hairs | and 13, more numerous comb scales, and a relatively long and

cylindrical rather than short and conical siphon. However, sierrensis and deserti-

cola both have larval hair 12-I present while it is absent in laguna, monticola and

varipalpus. In adult ornamentation, other than previously discussed, deserticola

and sierrensis show similarities, primarily in the reduced number of pleural setae

and smaller pleural scale patches, while laguna, monticola and varipalpus are alike

in having more pleural setae and more extensive pleural scale patches.

DISTRIBUTION. The Varipalpus Group is known only from western North

America from British Columbia near the southern border of Alaska to the Cape

Region of Baja California. The species with the most widespread distribution, sier-

rensis, extends from southern California and probably northern Baja California

through the Pacific drainage of California, Oregon, Washington, Idaho, Montana

and British Columbia and the margins of the interior basins of California, Nevada

and Utah. The remaining 4 species are represented by relict populations, laguna

in the mountains of the Cape Region of Baja California, monticola in southern

Arizona, varipalpus in northern Arizona and western Utah and deserticola in the

western margins of the Mohave and Colorado Deserts of southern California. All

species in the group are allopatric with the exception of deserticola and sierrensis

which occur together on the desert margins of the Transverse and Peninsular Ranges

of southern California.

AFFINITIES. The relationships of the Varipalpus Group to other mosquito

groups are obscure. It has been variously placed in several Aedes subgenera and

there is now considerable difference of opinion as to whether the group should

be placed in Finlaya or Ochlerotatus. Ludlow (1906:132) suggested that sierrensis

was probably a Finlaya and Dyar (1918:79, 1922:86) placed sierrensis (as vari-

palpus) in the subgenus Jaeniorhynchus on the basis of male genitalic characters

and Edwards (1932:154) placed varipalpus in group G of Finlaya, also on the

basis of male genitalia. Knight and Marks (1952:517) in their review of Finlaya

removed the Varipalpus Group from Finlaya and did not assign it to any other

subgenus, but Carpenter and LaCasse (1955:257) and Pratt (1956:10) placed it

in Finlaya. Belkin and McDonald (1957:190) removed the group from Finlaya

and placed it in Ochlerotatus ‘‘pending a much needed revision of this complex

subgenus and its various derivatives.”’ Rohlf (1963:802) in his numerical taxo-

nomic study of Aedes considered sierrensis to be a Finlaya as did Trebatoski

and Haynes (1969: 332) on the basis of enzyme comparisons, although the latter

included no species of Ochlerotatus in their study. Lunt and Nielsen (1971:110),

comparing adult thoracic setal characters, placed the Varipalpus Group in Group

X (Pulchritarsis-varipalpus Group) of Ochlerotatus and found no justification for

its placement in Finlaya. In his comparison of larval mouthparts, Gardner (1971:

80) suggested that the mandibles of the Varipalpus Group are ‘‘Finlayan” in form

and the maxillae closely resemble Aedes (Finlaya) togoi (Theobald, 1907), although

he did not actually place the group in Finlaya. It is our opinion that the Vari-

4 Contrib. Amer. Ent. Inst., vol. 8, no. 2, 1972

palpus Group fits in better in Ochlerotatus, as that complex subgenus is now de-

fined, than it does in Finlaya. It seems to show closer relationships to the Old

World Aedes (Ochlerotatus) pulchritarsis (Rondani, 1872) and related species than

to any other group (see discussion below). We feel, therefore, that it should be

retained in Ochlerotatus pending additional information on its affinities to other

groups.

Examination of other New World forms casts little light on the affinities of

the Varipalpus Group. A possible relationship between the Varipalpus Group and

Aedes muelleri Dyar, 1920 has been proposed, but the similarities are possibly

due to convergence. The male genitalia of muelleri are similar to those of laguna,

monticola and deserticola and although there are similarities between the larvae

and pupae of muelleri and those of the Varipalpus Group, some significant dif-

ferences are also present, such as the number and shape of the comb scales and

the development of the ventral brush. With the above mentioned exception of

the male genitalia, the adults of muelleri are unlike the Varipalpus Group, dif-

fering conspicuously in ornamentation, proboscis and leg length, female genitalia

and female tarsal claws. |

Belkin, Heinemann and Page (1970:143) briefly discuss a single male Aedes in

the USNM collection from Jamaica, probably an undescribed species. The printed

label reads //Jamaica BWI/Jan 1946/GA Thompson//. The abdomen of this spec-

imen is missing and there is apparently no genitalia slide extant. They place it

tentatively in the subgenus Ochlerotatus. The specimen shows some resemblance

to the Varipalpus Group in (1) decumbent scales of head curved, pointed, white

on vertex, dark laterally; (2) palpus with white over joints between segments 3

and 4 and 4 and 5 and few bristles on segments 3-5, but the palpus is consider-

ably shortened (0.69 of proboscis length instead of equal to proboscis); (3) meso-

notum with well-developed bristles and entirely covered with narrow, curved scales,

pale golden on dark brown background; (4) pleuron with patches of broad, white

appressed or semierect scales; (5) legs dark with pale scales forming ventral stripes

and basal and apical rings on femora, tibiae with basal and/or apical white scales

. (hindtibia missing) and tarsi with white scales basally and apically on segments

1 and 2; and (5) wing dark scaled. Since the genitalia are missing, it is impossible

to accurately place this specimen, but it may be allied to the Varipalpus Group.

There are several other treehole-breeding mosquitoes on the Island of Jamaica that

show relationships with the mosquito fauna of western North America. Haema-

gogus equinus Theobald, 1903 is found both in Jamaica and throughout Central

America as far north as northern Sinaloa in Mexico. Orthopodomyia waverleyi

(Grabham, 1907) of Jamaica is closely allied to O. signifera (Coquillett, 1896),

which is sympatric with the Varipalpus Group in Arizona and California. Coreth-

rella appendiculata Grabham, 1906 of Jamaica is found in western Mexico and the

U.S.A., as well as Central America and the Antilles south to Argentina. These

species may indicate a possible migratory pathway between Jamaica and Central

America at some time in the past which would account for a species allied to

the Varipalpus Group being present on Jamaica, although the evidence is very

tenuous and there are no other treehole Aedes or treehole Anopheles that show

this distribution pattern.

Aedes (Ochlerotatus) pulchritarsis (Rondani, 1872) and related species of the

Old World show many similarities to and may be the Old World representative

of the Varipalpus Group. Members of the pulchritarsis complex breed in tree-

holes and are found throughout the Mediterranean area, in central Asia and north-

Arnell & Nielsen: Aedes varipalpus Group 5

ern India and Pakistan. The male genitalia of pulchritarsis from Italy are very

much like those of varipalpus (sens. str.) with the exception of a poorly-developed

subapical lobe and variation in the distribution of setae on the dorsal surface of

the sidepiece. The male palpus is slightly longer than in the Varipalpus Group

and the terminal segments are more densely bristled. The species of this com-

plex diverge somewhat from the Varipalpus Group in details of mesonotal and

tarsal ornamentation, but otherwise the adults are similar. The larvae of the pul-

chritarsis complex have well-developed head hairs 5 and 6, longer antennae, comb

scales characterized by a long median spine, often an extremely long siphon and

5 pairs of hairs in the ventral brush. A thorough study of the chaetotaxy of the

immature stages of the pulchritarsis complex has never been undertaken. Since

the complex overall is so poorly known it is difficult to determine its affinities

but the many resemblances to the Varipalpus Group are suggestive of phyloge-

netic relationship.

SPECIATION AND PHYLOGENY. Based on the distribution patterns, rather

simple male genitalia and undoubtedly tropical origin, the Varipalpus Group ap-

pears to be an ancient one. It presents a mosaic of characters considering all stages

and consequently its phylogeny is difficult to sort out. Aedes laguna is prob-

ably closest to the ancestral stock. It has the most southerly distribution, the

male genitalia are the simplest with the undifferentiated seta of the basal lobe

more poorly developed and the larva is among the least specialized in the group.

The mesonotal ornamentation of 3 narrow lines of golden scales on the anterior

mesonotum is probably the primitive condition since this character is constant

in laguna and a tendency towards a reduction in golden scales is evident in the

remaining species of the group, all of which typically have a broad patch of gold-

en scales on the anterior mesonotum. Recent geological evidence (Anderson, 1971:

61) indicates that Baja California may have been connected to mainland Mexico

south and east of its present location as recently as 4-6 million years ago. Addi-

tional evidence for a relatively recent connection comes from biogeographical stud-

ies, a good example of which is in the distribution of Deinocerites mcdonaldi

Belkin & Hogue, 1959 found only on the Cape Region of Baja California and

mainland Mexico east and south of the Peninsula (Adames, 1971:81). If this is

the case, it would explain the existence of an isolated population of laguna on

Baja California. Proposals of migration southward along the Peninsula or disper-

sion westward to the Peninsula from the mainland across more than 100 miles

of open sea are untenable, and an alternate hypothesis of laguna being the relict

remnants of a more extensive population inhabiting a larger area of present-day

western Mexico can be proposed. Additional populations of laguna or a laguna-

like fauna might be expected in the higher elevations of the Sierra Madre Occi-

dental and western extremes of the Sierra Madre del Sur. Other species of tree-

hole mosquitoes have been taken in this area but it has not been sampled thoroughly.

Aedes monticola shows the closest affinity to laguna. The adult ornamentation

is similar except for the above-mentioned anterior mesonotal golden scales which

form a dense, broad patch in monticola. The male genitalia differ only in the

relative stoutness of the differentiated seta of the basal tergomesal lobe.

A light modification of the basal tergomesal lobe of the sidepiece of the male

genitalia of varipalpus indicates a more derived condition for this species. A char-

acteristic feature of varipalpus adults is a cluster of subspiracular setae. These setae

are also found in Jaguna although they are fewer in number and not always present.

Although the male genitalia of deserticola differ from laguna only in the rela-

6 Contrib. Amer. Ent. Inst., vol. 8, no. 2, 1972

tive stoutness of the differentiated seta of the basal tergomesal lobe and a ten-

dency toward fewer spines on the lobes on the ninth tergite, the larva and the

adult ornamentation differ to a greater extent. The larva of deserticola has hair

12-I developed and the setae and scales on the adult pleuron are much reduced

from the condition in laguna, monticola and varipalpus, indicating considerable

divergence from the more primitive members of the group. On the basis of the

male genitalia, deserticola was probably derived from laguna-monticola stock.

On the basis of its widespread distribution and relative abundance, as well as

apparently derived morphological characters, sierrensis would appear to be the

most modern and dominant species in the Varipalpus Group and probably diverged

from the remaining species comparatively recently. It exhibits the greatest amount

of intraspecific variation in the group. Based on very tenuous evidence of sim-

ilarities in adult ornamentation and the presence of hair 12-I in the larva and

present distribution patterns, it is possible that sierrensis is most closely allied to

deserticola. |

GENERAL CHARACTERISTICS

FEMALES. Small to medium-sized species; integument brown to black; dark

scales with slight iridescence; white markings on head, thorax, abdomen and legs,

and white and golden markings on mesonotum. Head: Eyes widely separated above

antennae. Decumbent scales white, pointed, curved, arranged on edge mesally and

on orbital border, continued forward between eyes; submedian patch of broad,

dark scales; orbital line expanded laterally into patch of broad, white appressed

scales, no orbital line below this patch; postgena with patch of broad, dark scales

with white scales below. Forked, erect scales over dorsal surface, yellow on ver-

tex, dark laterally. Two to 3 pairs of well-developed frontal bristles, usually mixed

light and dark; 3-5 pairs of well-developed dark orbital bristles. Clypeus dark brown

to black, bare. Proboscis short, 1.2-1.5 length of forefemur; entirely dark scaled

or with patch of white scales on basal ventral surface; a few basal bristles pres-

ent. Palpus about 0.25 of proboscis length; 4-segmented, segment 4 reduced, not

visible except in slide preparations; bristles on segments 1-3; dark scaled, occa-

sionally with white scales on apex of segment 1, apical dorsal surface of segment

2 and apical 0.25 of segment 3. Antenna slightly shorter than proboscis; torus

dark brown to black, with patch of white scales on mesal and dorsal surfaces;

flagellar segment 1 slightly longer than segment 2, with patch of white scales on

mesal surface; segments 2-13 with basal whorl of 6 bristles. Thorax: Mesonotum

moderately arched; mesonotal integument dark brown to black, lighter at scutal

angle and postalar margin; entire surface with scales except for normal anterior

inner dorsocentral and median and lateral prescutellar bare spaces; narrow, curved,

dark brown to black scales forming background with varied pattern of very nar-

row golden scales and broader yellow and white scales in lines or patches as fol-

lows: (1) median golden lines or broad patch from anterior promontory to near

prescutellar space, (2) narrow anterior crescent-shaped line formed from lateral

prescutal and posterior fossal lines, (3) narrow outer posterior dorsocentral line,

(4) rather broad supraalar line, (5) patches between outer posterior dorsocentral

line and supraalar line, and (6) anterior and lateral prescutellar lines. Acrostichal

bristles well developed, usually light anteriorly, dark posteriorly; dorsocentral bris-

tles longer than acrostichals, dark, merging posteriorly with dark prescutellars;

Arnell & Nielsen: Aedes varipalpus Group 7

humeral bristles strongly developed, mixed light and dark anteriorly; prescutal

bristles well developed, dark; usually 2 or 3 (1-5) dark posterior fossal bristles;

supraalar bristles numerous, well developed. Paratergite with numerous light or

dark bristles; cluster of white scales posteriorly. Scutellum dark brown; lobes with

patches of broad white scales and 4-8 long, dark bristles and several small, light

bristles. Postnotum dark brown. Pleural integument brown, lighter around spira-

cle and under wing base. Apn not enlarged. Pleural bristles on apn, ppn, ppl,

pst, psp, ssp (varipalpus and laguna), pra, stp, upper mep, lower mep (varipal-

pus and monticola). Apn with patch of mostly erect, narrow, curved, white scales;

ppn with large, rather sparse patch of narrow scales over entire sclerite, scales

black on anterodorsal border, white below (scales entirely white in varipalpus and

deserticola); patches of broad, white semierect or appressed scales on ppl, pst,

psp, long line on ssp, pra below knob, 2 patches on stp, 1 on upper margin and

1 posteriorly at level of midcoxa, the patches often joined by scattered scales,

patches on upper mep and lower mep, the patches sometimes continuous, pcx

(monticola and laguna), and small patch on metameron (except deserticola); hypo-

stigial area and meron without scales. Legs: Short, forefemur about 0.95-1.00 height

of thorax (mesonotum to apex of midcoxa). Hindcoxa slightly smaller than or

subequal to midcoxa, its base below upper margin of meron. Forecoxa with white

and brown scales, midcoxa and hindcoxa with white scales. Trochanters with

white scales. Femora mostly dark scaled, all with narrow basal and apical white

rings and scattered white scales; white scales forming streak on basal ventral sur-

face of forefemur and basal posterior surfaces of midfemur and hindfemur (an

additional basal anterior streak present on hindfemur of /aguna). Foretibia dark

scaled, with narrow basal and apical white rings; midtibia and hindtibia dark scaled,

with very narrow basal white rings and a few apical white scales. Foretarsus dark

scaled, with broad basal and apical white rings on segment 1, usually apical dor-

sal patch on segment 2; midtarsus as foretarsus except for scattered white scales

on apical portion of segment 5; hindtarsus dark scaled, with broad basal and api-

cal white rings on segment 1 (white scales occupying 0.4-0.5 of segment), broad

apical white rings on segments 2 and 3 (white scales occupying 0.3 of segment),

segment 4 with occasional basal and apical white scales except for sierrensis which

has broad basal white ring and scattered apical white scales, segment 5 entirely

white scaled. Foreclaws and midclaws subequal, slightly curved, with slender tooth

at basal third; hindclaws subequal, simple. Wing: Entirely dark scaled except for

white scales at base of C and occasionally base of R. Haltere: Yellowish, with

white scales on upper part of knob. Abdomen: Segments VIII and IX almost

completely retracted into VII. Tergite I with white and dark scales mesally, lat-

erotergite densely white scaled; tergites II-VI dark scaled, with narrow, curved

basal bands of white scales narrowed laterally and large basal rectangular white

patches on lateral margins; a few white scales apically on segment VI and often

V; tergite VII dark scaled, lacking basal white band but with lateral white patches

and rather broad apical white band. Sternites mostly white scaled, with variable

dark patches or bands becoming more extensive on distal segments. Cercus with-

out scales.

FEMALE GENITALIA. Segment VII: Tergite narrowed apically; length along

midline about 0.52-0.63 of tergite VII; uniformly covered with short bristles, a

few longer bristles present, more numerous distally. Sternite about 1.1-1.3 length

of tergite VIII, broader distally; uniformly covered with short bristles; a few long-

er bristles present, more numerous distally; very few scales occasionally present.

8 Contrib. Amer. Ent. Inst., vol. 8, no. 2, 1972

Tergite IX: Deeply emarginate apically; about 0.39-0.47 length of tergite VIII;

moderately sclerotized; apical lobes with 3-11 weakly developed bristles. Insula:

Moderately sclerotized, connected to sigma; with 1 or 2 pairs of moderately de-

veloped setae. Cercus: Moderately long, dorsal edge 0.65-0.82 length of tergite

VIII; compressed, apex rounded in lateral view; bristles numerous, the largest

strongly developed; scales absent. Postgenital Plate: Short and broad, 0.38-0.50

length of tergite VIII; index (length over width at midlength) about 1.30-1.50;

apex emarginate, distal portion with bristles, the longest moderately developed;

basal median longitudinal apodeme weakly to moderately sclerotized. Cowl: Strong-

ly sclerotized. Atrial plates absent. Sigma: Continuous with cowl, moderately scle-

rotized. Small portion of spermathecal duct strongly sclerotized. Spermathecae 3,

strongly sclerotized, spherical, 1 slightly larger than the others.

MALES. Generally similar to females except for sexual characters. Head: Pro-

boscis 1.3-1.6 length of forefemur, entirely dark scaled or with ventral white streak

on basal 0.4-0.5, often extending dorsally to form white ring from about basal

0.2-0.5. Maxillary palpus about equal to or slightly longer than proboscis; 5-seg-

mented; segment 1 dark scaled, with apical white band; segments 2 and 3 anky-

losed, making up 0.58-0.62 of palpus length, segment 3 expanded apically, both

segments without scales ventrally, remainder dark scaled with narrow white ring

at apex of segment 3, or with white scales over joint between segments 2 and

3 and white ring at apex of segment 3, subapical long bristles on segment 3; seg-

ments 4 and 5 turned down, dark scaled; segment 4 with narrow basal white

ring and numerous long, dark bristles; segment 5 with broad basal white ring and

long, dark bristles, more numerous apically. Antenna shorter than proboscis, torus

enlarged, with white scales on median and dorsal surfaces; flagellar segments 1-

12 with numerous long bristles; segment 1 elongate, with scales on mesal sur-

face; segments 12 and 13 elongate, the 2 combined slightly shorter than first 11

segments. Legs: Claws of foreleg and midleg enlarged, unequal; larger claws with

large, blunt subbasal tooth; smaller claws with acute subbasal tooth; claws of hind-

leg small, subequal, simple. Abdomen: Tergites II-VII lacking basolateral white

scale patches. Sternite VIII with large lateral white scale patches.

MALE GENITALIA. Tan to brown; apex of paraproct, midportion of tergite

IX and basal lobe of sidepiece darker. Segment IX: Tergite well developed; lobes

prominent, darkly pigmented, broader than long, separated by less than width of

1 lobe and bearing 2-10 strong setae. Sternite large, less than twice as broad as

long, with 3-8 setae apically. Sidepiece: Well developed, length 3.0-4.0 of width at

middle; lateral and ventral surfaces covered with strong bristles and scales; dor-

sal surface with short or strong bristles; tergomesal margin bare or with num-

erous short, strong hairs; apicotergal lobe absent; basal tergomesal lobe promi-

nent, with dense patch of strongly developed, apically curved setae or with sin-

gle strongly differentiated apically curved seta and 1 to many long, slender bris-

tles. Claspette: Stem moderately curved, length 4.0-8.0 of width, spiculate, with

several short setae; filament slightly longer than stem, recurved at apex; connec-

tion between claspettes declivous mesally or nearly horizontal with median emar-

gination. Clasper: Simple, moderately long (about 0.7 of sidepiece), slightly ex-

panded before middle, slightly to strongly curved apically; spiniform 0.25-0.30

length of clasper, slightly curved apically. Phallosome: Aedeagus cylindrical, length

2.0 of width, open ventrally. Proctiger: Strongly developed; basolateral scleroti-

zation vertical; paraproct well sclerotized, with a single large heavily sclerotized

apical tooth; cercal setae fine, short, 2-6.

Arnell & Nielsen: Aedes varipalpus Group 9

PUPAE. Cephalothorax: Yellowish to light brown, darker dorsally. Trumpet wid-

ened gradually from base to apex; amber to brown in color; tracheoid sculptur-

ing developed in basal 0.12-0.25, reticulate sculpturing well developed. Abdomen:

Weakly to darkly pigmented, darker anteriorly. Hair 1-II-[V weakly to moderately

developed, becoming weaker posteriorly, or subequal or becoming stronger on

segments II-I[V; usually double (1-3b) on II, single on III (1,2b), single on IV

(1,2b), weak and single on V and VI. Hair 2-III-VI mesad or laterad of hair 1

of corresponding segment. Hair 3 moderately to strongly developed on segments

IL III,V-VII; weakly developed, multiple on IV; at level or cephalad of 4-IV. Hair

5-IV-VI strongly developed, 0.6-1.0 length of corresponding segment; 5-VII longer

and stronger than 4-VII. Hair 7-V single to 4-branched. Hair 9 short and stout,

caudad of level of hair 6 and subequal in development on segments II-VI; strong-

ly developed, single (1,2b) on VII and considerably cephalad of caudolateral an-

gle of segment. Terminal Segments: Male genital lobe moderately large, about 1.4-

1.5 length of tergite VIII. Paddle: Length 1.3-1.8 of width. Apex emarginate or

occasionally rounded. Midrib conspicuous nearly to margin. Hair 1-P short, sin-

gle (1,2b).

LARVAE. Head: Width 1.2 of length. Lightly to moderately pigmented, lighter

in ocular area, darker posteriorly; collar deeply pigmented. Hair 1-C stout. Hairs

4,6-C displaced caudad, not near labrum. Hair 4-C moderately developed, 4-16b,

mesad of 1-C and at about same level as 6-C. Hair 5-C usually single (1,2b), slight-

ly mesad or laterad of 6-C. Hair 6-C usually double (1-3b). Hair 7-C 2-9b. Men-

tal plate with 8-12 teeth on each side of median tooth. Antenna: Length 0.4 of

head; shaft length usually 6.0-7.0 of basal width, with very few minute spicules.

Hair 1-A moderately long, inserted slightly beyond middle, single (1-3b). Tho-

rax: Integument pigmented or unpigmented, living larva white or gray in color;

without spicules. Tubercles of hairs 5-7-P separate. Hair 1-M,T moderately long,

branched. Hairs 4,5-P branched; 5-M single. Hair 11-P,M,T branched, much short-

er than 9-P. Hair 13-T moderately developed, usually 3-7b (2-10), subequal to

or slightly longer than 14-M. Hair 14-M moderately developed, usually 4,5b (2-

14). Abdomen: Integument as on thorax. Hair 1 moderately to strongly devel-

oped, usually increasing in development from segment I to V. Hair 2 cephalad

of level of hair 1 on segments I-VI, laterad of hair 1 on I,II,VI,VII and in line

with or mesad of hair 1 on IJJ-V. Hair 3-VII weakly developed. Hair 4 mesad

of hair 3 on segments III-V. Hair 6 usually double (1-4b) on segments I-V, sin-

gle or double on VI. Hair 7-I,II moderately to strongly developed, longer or short-

er than hair 10 of corresponding segment. Hair 12-I present or absent. Hair 13-

I,II moderately developed, 2,3b (2-6); hair 13-III-V strongly developed, usually

double (2,3b); hair 13-VI weakly developed, 3-15b, cephalolaterad of 10-VI. Seg-

ment VIII: Hair 1-VIII moderately to strongly developed, 1-4b (1-5); hair 3-VIII

strongly developed, 2-4b (1-5). Comb scales 7-20 (5-24), in irregular single to

triple row; individual comb scales elongate, more or less parallel-sided, with lat-

eral and apical fringe. Siphon: Strongly pigmented; relatively short to moderately

long (index 1.9-3.3); inflated at middle and reduced in diameter apically or with

sides nearly parallel (sierrensis); acus well developed. Pecten teeth 7-14 (7-19) in

number, restricted to basal 0.2-0.3 of siphon; each tooth with 1 or more sub-

basal spines. Hair 1-S strongly developed, usually triple (3-5b), 0.25-0.35 from

base of siphon. Anal Segment: Saddle moderate in size, extending about halfway

around segment; moderately to strongly pigmented, darker dorsally; imbricate

sculpturing indistinct. Hair 1-X strongly developed, single or double (1-3b). Hair

10

Contrib. Amer. Ent. Inst., vol. 8, no. 2, 1972

2-X triple or 4b. Hair 3-X single..Ventral brush (4-X) weakly developed, becom-

ing stronger distally, usually with 6 pairs of hairs, distal 5 pairs (4a-e-X) arising

from grid, usually double; proximal hair (4f-X) double to 6b. Gills sausage-shaped,

subequal; 3.0-5.5 of saddle length.

3(1).

4(3).

2(1).

3(2).

4(3).

KEYS TO SPECIES

FEMALES

Postcoxal area with. patch of white. scalese. 2s - Gecd cemee.ce ase darbeace e

Posteoxal.ates withteuG Scales.6 oo ho nl i ve Ee ee beh wh nD

Anterior part of mesonotum with 3 distinct narrow lines of golden scales

between dorsocentral bristles . . . . . . . 1. laguna

Anterior part of mesonotum with broad “patch of golden scales between

dorocentral Dristles:.. ¢. 25° ae oc a ie acwn 2. nontcola

Subspiracular area with several light setae arising from scale patch :

Siac ie eh eugea eaiet ld salt tiis 3. varipalpus

Subspiracular area without setae |

Base of hindtarsal segment 4 with broad white ring; metameron with patch

of white scales; anterodorsal border of ppn usually with dark scales.

. 5. sierrensis

Bae of fd eoemneat 4 Sper Oicad oie ae very narrow white

ring sometimes present; metameron without scales; ppn without dark

SCHR G ite ange Bok cro al tet nase hie einen. Gacetes GOSENTICOIE

MALE GENITALIA

Basal tergomesal lobe of sidepiece with dense patch of strongly developed,

apically curved setae; tergomesal margin of sidepiece with numerous short,

Sirone hairs... ... . . 5. Sierrensis

Basal tergomesal lobe of sidepiece with ia ong: differentiated, apically

curved seta and 1 to several slender bristles; tergomesal margin of side-

poe OU er Onan ny 19

Basal tergomesal lobe of sidepiece with 4 or more long slender bristles in

addition to strongly differentiated seta . . . . 3. varipalpus

Basal tergomesal lobe of sidepiece with 1-3 long slender bristles in addi-

Hom to. stronely dittcrentizied seta ~.. "as. 3 ot ee ke

Differentiated seta of basal tergomesal lobe about 40 times as long as its

Dasai width... . . . . 1. laguna

Differentiated seta of basal tergomesal lobe about 5 30 times as long as

POS eee ee ae eee oh ge a

Lobes of tergite [IX usually with 6 or 7 (4-10) setae .. . . 2. monticola

Lobes of tergite [X usually with 4-6 (2-7) setae . . . . . . 4. deserticola

2(1).

3(1).

4(3).

aT).

3(2).

4(3).

Arnell & Nielsen: Aedes varipalpus Group 11

PUPAE

Abdominal segments I-V pigmented brown at least mesally. . . . . . .2

Abdominal segments I-V pigmented light yellowish-brown. . . . .. .3

Entire abdomen pigmented brown except for light yellowish-brown areas

on lateral margins of segments V-VII]. ... . ei i laguna

Abdominal segment I and median anterior portions ‘of segments II-IV pig-

mented brown, remainder of abdomen light yellowish-brown .

. 5. sierrensis

Hair 1-IV more strongly developed than 1-1 or 1-V and usually double.

: 3. varipalpus

Hair 1IV subequal in development ‘to 1-Ill and eae :

Cephalothorax light yellowish-brown with darker pigmentation on middorsal

line; hair 1-II usually single (1-3b) . . . . 2. monticola

Cephalothorax Hae ee A yellowish-brown: hair ac usually double

2) Breer & j ragga . . « « « « 4, deserticola

LARVAE

Body pigmented, gray in color; siphon relatively long (index usually at

least 3.0), not inflated at middle, not reduced in diameter apically; comb

scales usually more than 15... . . . 3. sierrensis

Body unpigmented, white in color; siphon short @ndex less than 2.8), in-

flated near middle and sharply reduced in diameter apically; comb scales

Ustially Tess Chiaie a5 ea ag OR Ie ere ny gee enc gt ae

Hair 1-X usually single, rarely double; 1-VIII frequently single; pecten teeth

few in number (7-11) and restricted to basal 0.20 of siphon . ie

ae . 4. deserticola

Hair 13% double, "rarely ‘single: ‘1-VIII usually ‘triple, never single; pecten

teeth more numerous (10-14) and extending to at least basal 0.25 of

SHON ORAS. CA: BREN! ROLE Sau ARE, FPS ARNE ARR EO ea SR Oe

Siphon index 2.6-2.8 (2.5-3.0); hair 6-VI usually double. . . . . 1. laguna

Siphon indéx dess*than 2:5 hair 6-VF single) 2) eV a er,

Hairs 1-IV,V usually with same number of branches and frequently sub-

equal to hair 13 of corresponding segment; 7-P frequently triple; anal

gills about 3.5 of saddle length . . . . 2. monticola

Hairs 1-IV,V usually with more branches than and usually weaker than

hair 13 of corresponding segment; 7-P usually double, rarely triple; anal

gills usually about 4.5 of saddle length. . . ... . . . 3. varipalpus

12 Contrib. Amer. Ent. Inst., vol. 8, no. 2, 1972

TAXONOMIC TREATMENT

1. Aedes (Ochlerotatus) laguna Arnell & Nielsen, n.sp.

Figs. 1-4

TYPES: Holotype male (UCLA 695-12) with associated larval and pupal skins and genitalia

slide, La Laguna meadow, elevation 6800 ft, Sierra Laguna, 20 mi ENE Todas Santos, Baja

California Sur, Mexico, larva from a rothole in a living oak (Quercus sp.), 27 Aug 1970, J.H.

Arnell and L.T. Nielsen [USNM]. Allotype female (UCLA 695-21) with associated larval and

pupal skins, same data as holotype [USNM]. Paratypes: 3 lpF (UCLA 695-11,17,18), 39 M,

60 F, 98 P, 239 L (UCLA 695), same data as holotype [UCLA, UTAH, USNM].

Aedes (Ochlerotatus) monticola of Belkin and McDonald (1957:188).

Aedes (Finlaya) varipalpus of Ross (1943:86).

FEMALE (fig. 2). Wing: 3.65 mm. Proboscis: 2.55 mm. Forefemur: 1.95 mm.

Abdomen: about 2.70 mm. Head: Erect scales mostly yellow; a few dark erect

scales laterally. Proboscis dark scaled, with small patch of white scales on basal

ventral surface. Palpus with segment 1 entirely dark scaled. Thorax: Mesonotum

with narrow line of narrow golden scales along acrostichal line from anterior pro-

montory to near prescutellar space; stripe of golden scales just mesad of anter-

ior dorsocentral line from prescutal area to middle of mesonotum; outer poster-

ior dorsocentral line composed of yellow and white scales; patches between outer

dorsocentral line and supraalar line reduced; anterior prescutellar patch white.

Acrostichal bristles mostly light, a few dark. Pon with narrow dark scales anter-

iorly and broader white scales in sparse patch posteriorly, scales becoming more

dense posteroventrally, 4-8 light setae posteriorly; ppl with about 30 light setae;

ssp often with 1-3 light setae arising from scale patch; pcx with patch of white

scales. Foretarsal segment 2 with apical dorsal white scale patch reduced or ab-

sent; midtarsal segment 5 entirely dark scaled; hindfemur with anterior and pos-

terior basal white streak; hindtarsal segment 1 with broad basal and apical white

rings, white scales occupying about 0.5 of segment; hindtarsal segment 4 with

very narrow apical white ring.

MALE. Wing: 3.00 mm. Proboscis: 2.80 mm. Forefemur: 1.80 mm. Abdomen:

about 2.90 mm. Proboscis with ventral white streak on basal 0.4, extending dor-

sally to form ring from about basal 0.2-0.4: Palpus without white scales over joint

between segments 2 and 3. or

MALE GENITALIA (fig. 3). Segment IX: Tergite lobe with about 6 (48) se-

tae. Sidepiece: Length 4.0 of width at middle; lateral 0.33 of dorsal surface with

strong bristles and scales, mesal 0.67 of dorsal surface with short bristles becom-

ing stronger toward apex; tergomesal margin bare; basal tergomesal lobe with sin-

gle strongly differentiated, apically curved seta and 1-3 long slender bristles; length

of differentiated seta about 40.0 of width at base. Claspette: Stem moderately

curved, length 8.0 of width; filament slightly longer than stem, flattened beyond

middle, sharply recurved at apex; connection between claspettes declivous me-

sally. Clasper: Strongly curved apically; spiniform about 0.3 of clasper length.

PUPA (fig. 3). Abdomen: 3.35 mm. Trumpet: 0.51 mm. Paddle: 0.82 mm.

Cephalothorax: Light brown, darker dorsally; trumpet dark yellowish-brown; meta-

notum brown, haltere case lighter. Abdomen: Dark brown, yellowish on lateral

margins and posterior segments. Hair 1-II usually double, often triple, rarely sin-

gle; 1-III-V subequal in development. Hair 5-III subequal to 1-III and weaker than

Arnell & Nielsen: Aedes varipalpus Group 13

3-III. Hairs 5-IV-VI strongly developed, subequal. Hair 7-V usually single. Pad-

dle: Apex emarginate; length 1.3-1.5 of width.

LARVA (fig. 4). Head: 0.83 mm. Siphon: 0.89 mm. Saddle: 0.28 mm. Tho-

rax: Integument unpigmented, living larva white. Hair 7-P usually triple, rarely

double. Abdomen: Integument as on thorax. Hair 1 stronger than hair 13 on

segment II, generally weaker and with more branches than hair 13 on III-V. Hair

6 usually double on segments I-VI. Hair 7-I,II well developed, stronger than hair

10 on corresponding segment. Hair 12-I absent. Segment VIII: Hair 1 usually

triple (2-4b); hair 3 usually triple (2,3b). Comb scales 12-15 (10-22), in single

or irregular double row. Siphon: Index 2.6-2.8 (2.5-3.0); inflated just before mid-

dle, sharply reduced in diameter apically. Pecten teeth 11-14 (10-17); restricted

to basal 0.25 of siphon. Hair 1-S triple, 0.25-0.35 from base of siphon. Anal

Segment: Hair 1-X usually double (1-3b). Gills about 4.0 of siphon length.

SYSTEMATICS. The pattern of golden scales on the mesonotum of adults of

laguna is distinctive, there being 3 discrete narrow lines on the anterior part rather

than a broad patch characteristic of the other members of the group. One to 3

light setae arise from the subspiracular scale patch in many adults and the post-

coxal scale patch is always present. The male genitalia of laguna, although very

similar to those of monticola and deserticola, with 1 differentiated seta and 3

or fewer stout bristles on the basal tergomesal lobe of the sidepiece, can be dis-

tinguished from the latter 2 species by the differentiated seta being relatively fine

and its length about 40 times its basal diameter instead of 25-30 times as in mont-

icola and deserticola. The brown pigmentation of the cephalothorax and abdo-

men of the pupa of laguna is unique among the Varipalpus Group and there

are slight differences in chaetotaxy from the other species, hair 1-II being usu-

ally double and 7-V usually single. The larva of laguna can be distinguished from

the other members of the group primarily by the more elongate (index 2.5-3.0)

and medially inflated siphon, hair 6-VI usually double and 7-P usually triple.

Aedes laguna is probably close to the parental stock which gave rise to the

Varipalpus Group and may be a relict remnant of a more extensive population

that inhabited western Mexico prior to the separation of Baja California from the

mainland. A more thorough discussion is presented under group systematics above.

BIONOMICS. Aedes laguna has been collected at only 1 locality, a meadow

at an elevation of about 6,800 feet near the top of the Sierra Laguna, approxi-

mately 20 miles ENE of Todos Santos, Baja California Sur, Mexico. The area is

a cloud forest of pine and oak with a ground cover of grasses and low shrubs.

Larvae of laguna were collected from 2 treeholes in living oak (Quercus sp.). One

hole was small, containing less than 50 ml of water and few larvae. The other

hole was approximately 8 inches in diameter, 6-8 inches deep and contained about

2 liters of water. Several hundred larvae were removed, the majority of them in

the fourth instar. No eggs were obtained from the treeholes in the area. Several

collections of Orthopodomyia kummi Edwards, 1939 were made in treeholes in

the same area but these did not contain laguna. A mermithid nematode parasite

was found in about 25 laguna larvae. The nematode was usually aligned longi-

tudinally in the haemocoele of the fourth instar larva and coiled back on itself

once. It was observed to exit the mosquito body through the siphon, leaving the

larva dead or dying. Nothing is known of the habits of laguna except that fe-

males are avid biters. Within 5 minutes, at noon with low clouds and light mist,

about 15 females were captured while attempting to bite and many more avoided

capture.

14 Contrib. Amer. Ent. Inst., vol. 8, no. 2, 1972

_ DISTRIBUTION (fig. 1). Mountains of the Cape Region of Baja California. All

material examined is listed above in the type series.

2. Aedes (Ochlerotatus) monticola Belkin & McDonald

Figs. 1,2,5,6

1957. Aedes (Ochlerotatus) monticola Belkin and McDonald, 1957:179-191. TYPE: Holotype

male (UCLA 173-104) with associated larval and pupal skins and genitalia slide, Madera

Canyon (Santa Cruz Co.), Arizona, U.S.A., elevation ca 5000 ft, 18 Aug 1955, W.A.

McDonald [USNM, 63244].

Aedes (Ochlerotatus) monticola of McDonald (1957:505); Zavortink (1970:1); Lunt and Nielsen

(1971:110).

Aedes monticola of Nielsen, Arnell and Linam (1967:74); Nielsen, Linam, Arnell and Zavortink

(1968:362).

Aedes (Finlaya) varipalpus in part of Carpenter and LaCasse (1955:257).

Aedes varipalpus in part of Jenkins and Carpenter (1946:39); Peyton (1956:220); Richards,

Nielsen and Rees (1956:14).

Aedes (Taeniorhynchus) varipalpus in part of Dyar (1922:86).

FEMALE (fig. 2). Wing: 3.45 mm. Proboscis: 2.20 mm. Forefemur: 1.75 mm.

Abdomen: about 2.65 mm. Head: Erect scales yellow on vertex; numerous dark

erect scales laterally. Proboscis dark scaled, with small patch of white scales on

basal ventral surface. Palpus with a few white scales on apex of segment I. Tho-

rax: Mesonotum with broad dense patch of narrow golden scales from anterior

promontory to near prescutellar space, often narrow lines of white scales anter-

iorly at lateral margin of golden patch and anteriorly along acrostichal line; lat-

eral prescutal white scales relatively sparse; outer posterior dorsocentral line broad

anteriorly, composed of white and golden scales, narrowed and composed of white

scales posteriorly; supraalar line narrowed; patches between outer posterior dor-

socentral line and supraalar line conspicuous, golden anteriorly and white poster-

iorly; anterior prescutellar patch white. Acrostichal bristles dark. Pon with nar-

row dark scales anteriorly and broader white scales in sparse patch posteroven-

trally, 4-10 mixed light and dark setae posteriorly; pp/ with about 30 light setae;

ssp without setae; pcx with dense tuft of white scales; lower mep usually with

2-4 light setae arising from cephalodorsal margin of scale patch, scale patch large,

often continuous with scale patch on upper mep. Hindtarsal segment 1 with broad

basal and apical white rings, white scales occupying about 0.5 of segment; hind-

tarsal segment 4 entirely dark scaled.

MALE. Wing: 2.75 mm. Proboscis: 2.20 mm. Forefemur: 1.65 mm. Abdomen:

about 2.75 mm. Proboscis with ventral white streak on basal 0.5, usually extend-

ing dorsally to form white ring from about basal 0.3-0.5. Palpus without white

scales over joint between segments 2 and 3.

MALE GENITALIA (fig. 5). Segment IX: Tergite lobe bearing 6 or 7 (4-10)

setae. Sidepiece: Length 4.0 times width at middle; lateral 0.33 of dorsal surface

with strong bristles and scales, mesal 0.67 of dorsal surface with short bristles

becoming stronger toward apex; tergomesal margin bare; basal tergomesal lobe

with single strongly differentiated, apically curved seta and 1-3 long, slender bris-

tles; length of differentiated seta about 25.0-30.0 of width at base. Claspette:

Stem moderately curved, length 8.0 of width; filament slightly longer than stem,

Arnell & Nielsen: Aedes varipalpus Group 15

flattened beyond middle, sharply recurved at apex; connection between claspettes

declivous mesally. Clasper: Strongly curved apically; spiniform about 0.3 of clasper

length.

PUPA (fig. 5). Abdomen: 3.78 mm. Trumpet: 0.47 mm. Paddle: 0.89 mm.

Cephalothorax: Light yellowish-brown, darker in caudal middorsal area; trumpet

amber; metanotum darker mesally. Abdomen: Light yellowish-brown, slightly dark-

er mesally on anterior segments. Hair 1-II usually single; 1-III-V subequal in de-

velopment. Hair 5-III usually subequal to 1-III and more weakly developed than

3-III but often subequal to 3-III and stronger than 1-III. Hairs 5-IV-VI strongly

developed, subequal. Hair 7-V usually double. Paddle: Apex emarginate, length

1.3-1.6 of width.

LARVA (fig. 6). Head: 0.96 mm. Siphon: 0.90 mm. Saddle: 0.30 mm. Tho-

rax: Integument unpigmented, living larva white. Hair 7-P double or triple. Ab-

domen: Integument as on thorax. Hair 1 stronger than hair 13 on segment II,

weaker and with more branches than hair 13 on III and often subequal to hair

13 (2b) on IV and V. Hair 6 usually double on segments I-V, single on VI. Hair

7-I,I] well developed, stronger than hair 10 on corresponding segment. Hair 12-I

absent. Segment VIII: Hairs 1 and 3 usually triple (2,3b). Comb scales 9-13 (7-

17), in single or irregular double row. Siphon: Index 2.2-2.5; concave on basal

ventral surface, inflated just beyond middle, sharply reduced in diameter apically.

Pecten teeth 11-14 (8-17); restricted to basal 0.3 of siphon. Hair 1-S usually trip-

le (3-5b), 0.30-0.35 from base of siphon. Anal Segment: Hair 1-X usually double

(2-4b). Gills usually about 3.5 of saddle length.

SYSTEMATICS. The mesonotal scale pattern of adult monticola is more sharply

defined than in the other members of the group, the golden and white scales

being more densely clustered and prominent, with no intermediate yellow scales.

The pleural setae are more numerous and the pleural scale patches are larger with

the scales more erect, lending a shaggy appearance to the pleuron. Present are

dense clusters of scales in the postcoxal area and lower mesepimeral setae. The

male genitalia of monticola are almost indistinguishable from those of laguna and

deserticola with 1 differentiated seta and 3 or fewer stout bristles on the basal

tergomesal lobe of the sidepiece. However the differentiated seta is stouter in

monticola than in laguna and the lobes of the ninth tergite generally have more

setae (6 or 7) in monticola than in deserticola. The pupa of monticola is sim-

ilar in pigmentation to varipalpus but can be distinguished from it by hairs 1-

III-V being subequal. The larva of monticola is readily differentiated from laguna

and deserticola by the characters given in the key, but its separation from vari-

palpus by the key characters is not always reliable. The siphon of monticola is

usually longer than in varipalpus, with an index of 2.2-2.5 in the former and 1.9-

2.4 in the latter.

Hybridization between monticola and varipalpus is apparent in specimens col-

lected at 2 localities in Oak Creek Canyon, Arizona, and is discussed under vari-

palpus.

BIONOMICS. The adults of monticola attack man readily but nothing else is

known of their habits. Larvae have been collected from treeholes in oak (Quercus

sp.), cottonwood (Populus fremontii), willow (Salix sp.) and sycamore (Platanus

racemosa). This species is found primarily in riverine situations in cottonwood,

willow and sycamore over much of its range but is widespread in the oak wood-

land at the higher elevations in southeastern Arizona and adjacent New Mexico

and Sonora.

16 Contrib. Amer. Ent. Inst., vol. 8, no. 2, 1972

Aedes monticola has been found with varipalpus at 1 locality in Oak Creek

Canyon; with Toxorhynchites sp. in 2 collections in Cochise and Santa Cruz Coun-

ties; with Aedes purpureipes Aitken, 1941 and an undescribed species in the Aedes

kompi group from several localities in Pima and Santa Cruz Counties and | lo-

cality in Sonora; with Orthopodomyia kummi Edwards, 1939 in southeastern Ari-

zona, and with Aedes muelleri Dyar, 1920, Anopheles judithae Zavortink, 1969

and Orthopodomyia signifera (Coquillett, 1896) throughout most of its range. In

1 willow treehole in Hidalgo County, New Mexico, monticola larvae were found

in association with muelleri, judithae, signifera and kummi. Under laboratory con-

ditions, at temperatures of 65° to 75° F, monticola developed from eggs to adults

in 23 days.

DISTRIBUTION (fig. 1). Aedes monticola has been collected in the Gila River

drainage of Arizona and New Mexico, 1 collection in the Little Colorado River

drainage of Arizona and 1 collection in the Rio Magdalena drainage of Sonora.

Material examined: 1897 specimens; 325 males, 346 females, 801 larvae, 425 pu-

pae; 206 individual rearings (142 larval, 64 pupal).

MEXICO

Sonora. Nogales, 8.8 mi S Mex.-U.S. border, 13 June 1971, TJZ, LTN (MEX 703), 2 lpM

(703-10,12), 3 IpF (703-11,13,14), 1 M, 3 P [UCLA].

UNITED STATES

Arizona. Cochise Co.: Carr Canyon, 5 Sept 1966, TJZ (UCLA 338), 1 lpM (338-10) [UCLA] ;

same data (UCLA 339), 2 IpM (339-11,19), 6 IpF (339-10,12,13,15-17), 10 pM (339-40,41,100-

107), 4 pF (339-108-111), 5 M, 10 P, 5 L [UCLA]. Cave Creek Canyon, 15 June 1963, J.

Burger (UCLA 387), 1 F [UCLA]. Cochise Stronghold, Dragoon Mts., 5000 ft, 26 Aug 1964, J.

Burger (UCLA 257), 9 F [UCLA]; same data, 22 Mar 1966, TJZ (UCLA 302), 1 IpM (302-16)

[UCLA]; same data (UCLA 303), 5 IpM (303-10,13,14,17,40), 6 IpF (303-11,12,15,16,18,19),

1 pM (303-100), 1 pF (303-101), 2 P, 5 L [UCLA]; same data (UCLA 304), 3 IpM (304-10,17,

42), 11 pF (304-11-16,18,19,41,43), 12 pM (304-40,100-109,113,114), 4 pF (304-110-112,114),

18 M, 6 F, 25 P, 48 L [UCLA]; same data (UCLA 305), 6 lpM (305-10-13,17,18), 4 IpF (305-

14-16,19), 2 pM (305-100,101), 1 pF (305-102), 3 L [UCLA]; same data (UCLA 306), 4 IpM

(306-10-12,18), 6 IpF (306-13-17,19), 16 pM (306-41,43,45,100-112), 3 pF (306-40,113,114),

1M, 2 F,4P, 3 L [UCLA]; same data (UCLA 427), 1 F, 1 P, 1 L [UCLA]; same data, 4 Sept 1966

(UCLA 325), 1 pF (325-100) [UCLA]; same data (UCLA 326), 1 M, 2 F,4P, 1 L [UCLA];

same data (UCLA 329), 4 IpF (329-10-13), 5 M, 13 F, 27 P, 5 L [UCLA]; same data (UCLA

330), 1 IpF (330-10), 1 pF (330-111) [UCLA]; same data (UCLA 332), 1 F [UCLA]. Coronado

Natl. Mon., 5200 ft, 20 Mar 1968, HA, LTN, JHL (UCLA 659), 2 M, 4 F [UCLA-UTAH].

Southwest Research Station, 5 mi SW Portal, 5-25 Sept 1965, C.W. Sabrosky, 1 F [USNM].

Coconino Co.: Oak Creek Canyon, Manzanita Campground, 4700 ft, 24 May 1970, HA, LTN,

RN (UCLA 682), 1 IpM (682-22), 5 IpF (682-11,14,15,20,23), 11 M, 7 F, 7 L [UCLA-UTAH].

Gila Co.: Armer Mtn. Trail, Sierra Ancha Mts., 25 May 1963, J. Burger (UCLA 381), 3 F [UCLA].

Globe, 3541 ft, 22 Mar 1966, LTN (UCLA 636), 19 M, 22 F [UCLA-UTAH]. Globe (19 mi

NE), 4500 ft, 2 Sept 1966, TJZ (UCLA 331), 1 F, 2 P [UCLA]. Navajo Co.: Mortensen Wash, 14

mi W Show Low, 14 Aug 1966, LTN (UCLA 637), 5 M, 6 F [UCLA-UTAH]. Pima Co.: Babo-

quivari Camp, Baboquivari Mts., 3550 ft, 31 Dec 1969, TJZ (UCLA 558), 1 lpM (558-32) [UCLA].

Sabino Canyon, Baboquivari Mts., 3900 ft, 30 Dec 1969, TJZ (UCLA 554), 5 lpM (554-20,21,

24-26), 5 IpF (544-22,23,29,34,35), 1 L [UCLA]; same data (UCLA 560), 3 IpM (560-30-32),

1 IpF (560-33), 10 L [UCLA]. Sycamore Canyon, Baboquivari Mts., 3500-3700 ft, 31 Dec

1969, TJZ (UCLA 557), 1 IpM (557-10) [UCLA]; same data (UCLA 567), 1 M, 1 P,1L

[UCLA]. Mendoza Canyon, Coyote Mts., 3500 ft, 7 Sept 1969, TJZ (UCLA 627), 6 F [UCLA];

same data, 29 Dec 1969, TJZ, LTN (UCLA 551), 2 pM (551-100,101) [UCLA]; same data

(UCLA 552), 1 IpM (552-21), 4 IpF (552-20,22-24), 1 M, 1 F, 2 P, 14 L [UCLA]; same data

(UCLA 555), 1 IpF (555-31) [UCLA]; same data (UCLA 561), 1 IpM (561-30), 3 IpF (561-

31-33) [UCLA]; same data (UCLA 653), 3 IpF (653-15,17,18), 1 M, 5 F, 24 L [UCLA-UTAH] ;

Arnell & Nielsen: Aedes varipalpus Group 17

same data (UCLA 654), 4 M, 13 F [UCLA-UTAH]; same data, 30 Dec 1969 (UCLA 553), 4 IpM

(553-21,23-25), 8 IpF (553-20,22,26-31), 82 M, 48 F, 122 P, 467 L [UCLA]. Molino Basin,

Santa Catalina Mts., 5000 ft, 2 June 1963, J. Burger (UCLA 385), 1 F [UCLA]. Pinal Co.:

Aravaipa Canyon, 11 mi E Hwy. 77, 2600 ft, 21 Dec 1966, LTN (UCLA 638), 2 M, 2 F, 1 P

3 L [UCLA-UTAH]. Santa Cruz Co.: Calabasas Canyon, 21 Mar 1966, TJZ (UCLA 296), 1 IpM

(296-10), 2 lpF (296-11,12), 2 pM (296-100,102), 1 P, 1 L [UCLA]; same data (UCLA 297),

3 IpM (297-10,11,13), 6 lpF (297-12,14-18), 4 pM (297-100-103) [UCLA]; same data, 13 Sept

1968 (UCLA 451), 3 pM (451-10,100,101), 1 pF (451-102) [UCLA]. Lochiel (11 mi E), 5100

ft, 21 Mar 1968, HA, LTN, JHL (UCLA 660), 3 M, 5 F, 3 L [UCLA-UTAH] ; same data, 27 Dec

1969, TJZ, LTN (UCLA 559), 1 L [UCLA]; same data (UCLA 562), 1 L [UCLA]. Madera

Canyon, Santa Rita Mts., 5000 ft, 17 Aug 1955, W.A. McDonald (UCLA 170), 12 F [UCLA];

same data, 18 Aug 1955 (UCLA 173), 2 IpF (173-102,103), 1 P, 2 L [UCLA]; same data,

30 June 1956 (UCLA 206), 2 L, E [UCLA]; same data (UCLA 219), 1 lpM (219-103) [UCLA] ;

same data, 21 Aug 1970, HA, LTN (UCLA 694), 7 IpF (694-11-17), 37 M, 52 F, 19 L [UCLA-

UTAH]. Patagonia (1.7 mi SW), 4000 ft, 21 Mar 1968, HA, LTN, JHL (UCLA 662), 4 M, 4 F

[UCLA-UTAH]; same data, 13 Sept 1968, TJZ (UCLA 455), 1 lpM (455-10) [UCLA]; same

data (UCLA 460), 1 IpM (460-10), 2 IpF (460-12,16); same data, 14 Sept 1968 (UCLA 463),

1 IpM (463-10), 1 lpF (463-11) [UCLA]; same data, 14 Sept 1968 (UCLA 471), 1 IpF (471-

15) [UCLA]; same data (UCLA 472), 1 lpM (472-20) [UCLA]. Yavapai Co.: Prescott (7.5 mi

NNE), 5150 ft, 19 Mar 1966, LTN (UCLA 635), 3 M, 4 F, 1 P, 3 L [UCLA-UTAH]. Wet

Beaver Creek, 2 mi E I-17, 3700 ft, 29 Aug 1970, HA, LTN (UCLA 696), 10 M, 8 F, 8 P,

29 L [UCLA-UTAH]. Verde River at Hwy. 79, 3200 ft, 22 Mar 1968, HA, LTN (UCLA 663),

1 F [UCLA].

New Mexico. Hidalgo Co.: Animas (15 mi S), 4700 ft, 19 Mar 1968, HA, LTN, JHL (UCLA

657), 34 M, 44 F [UCLA-UTAH].

3. Aedes (Ochlerotatus) varipalpus (Coquillett)

Figs. 1,2,7-9

1902. Culex varipalpus Coquillett, 1902:292-293. TYPE: Holotype female, Williams (Coconino

Co.), Arizona, U.S.A., 29 July 1902, H.S. Barber [USNM, 6559].

Aedes (Ochlerotatus) varipalpus of Belkin and McDonald (1957:179,189,191); Myers (1967:806);

Zavortink (1970:1); Lunt and Nielsen (1971:110).

Aedes varipalpus of Dyar and Knab (1906:200); McDonald (1957:505); Nielsen and Rees (1959:

46); Nielsen, Arnell and Linam (1967:75).

Aedes varipalpus in part of Howard, Dyar and Knab (1917:644); Jenkins and Carpenter (1946:

39); Peyton (1956:220); Richards, Nielsen and Rees (1956:14).

Aedes (Finlaya) varipalpus of Nielsen and Rees (1961:33).

Aedes (Finlaya) varipalpus in part of Edwards (1932:154); Carpenter and LaCasse (1955:257);

Pratt (1956:6).

Aedes (Taeniorhynchus) varipalpus in part of Dyar (1922:86).

Aedes (?) varipalpus of Knight and Marks (1952:517).

Ochlerotatus varipalpus of Coquillett (1906:20).

Culex varipalpus of Blanchard (1905:628,630); Theobald (1910:365).

FEMALE (figs. 2,7). Wing: 3.68 mm. Proboscis: 2.40 mm. Forefemur: 1.85

mm. Abdomen: about 2.80 mm. Head: Erect scales mostly yellow; a few dark

erect scales laterally. Proboscis dark scaled, with small patch of white scales on

basal ventral surface. Palpus with a few white scales on apex of segment 1. Tho-

rax: Mesonotum with broad patch of narrow golden scales from anterior pro-

montory to about middle of mesonotum, this patch sparse, blending gradually

into area of dark scales laterally; outer posterior dorsocentral line of yellow and

18 Contrib. Amer. Ent. Inst., vol. 8, no. 2, 1972

white scales; patches between outer posterior dorsocentral line and supraalar line

golden and yellow, scales sparse; anterior prescutellar patch mixed yellow and

white. Acrostichal bristles dark. Ppn with large, sparse patch of white scales over

entire sclerite, more numerous posteroventrally, single or double row of 6-12 mix-

ed light and dark setae posteriorly, usually 2 or 3 light setae continuing toward

ventral margin of sclerite; ppl with about 30 light setae; ssp with 4-6 (2-9) light

setae arising from scale patch; pcx without scales; lower mep usually with 2-4

light setae arising from cephalodorsal margin of scale patch. Hindtarsal segment

1 with broad basal and apical white rings, white scales occupying 0.5 of segment;

hindtarsal segment 4 usually with small basal dorsal patch of white scales, re-

mainder dark scaled.

MALE (fig. 7). Wing: 3.05 mm. Proboscis: 2.35 mm. Forefemur: 1.70 mm.

Abdomen: about 2.75 mm. Proboscis with ventral white streak on basal 0.5, us-

ually extending dorsally to form white ring from about basal 0.3-0.5. Palpus with- _

out white scales over joint between segments 2 and 3.

MALE GENITALIA (fig. 8). Segment [X: Tergite lobe with 5 or 6 (3-10) se-

tae. Sidepiece: Length 4.0 of width at middle; lateral 0.33 of dorsal surface with

strong bristles and scales, mesal 0.67 of dorsal surface with short bristles becom-

ing stronger toward apex; tergomesal margin bare; basal tergomesal lobe with sin-

gle strongly differentiated, apically curved seta and 4 or more long, slender bris-

tles. Claspette: Stem moderately curved, length 8.0 of width; filament slightly

longer than stem, flattened beyond middle, sharply recurved at apex; connection

between claspettes declivous mesally. Clasper: Strongly curved apically; spiniform

about 0.3 of clasper length.

PUPA (fig. 8). Abdomen: 3.89 mm. Trumpet: 0.56 mm. Paddle: 0.79 mm.

Cephalothorax: Light yellowish-brown, darker on caudal middorsal area; trumpet

amber; metanotum darker mesally. Abdomen: Light yellowish-brown, slightly dark-

er mesally on anterior segments. Hair 1-II usually single; 1-IV relatively strongly

developed, usually double (1-3b), longer than 1-III or 1-V. Hair 5-III usually sub-

equal to 1-III and more weakly developed than 3-III but often subequal to 3-

III and stronger than 1-IIJ. Hairs 5-IV-VI strongly developed, subequal. Hair 7-V

double. Paddle: Apex emarginate; length 1.4-1.6 of width.

LARVA (fig. 9). Head: 0.78 mm. Siphon: 0.60 mm. Saddle: 0.19 mm. Thorax:

Integument unpigmented, living larva white. Hair 7-P double or triple. Abdomen:

Integument as on thorax. Hair 1 stronger than hair 13 on segment II, weaker

and usually with more branches than hair 13 on III-V. Hair 6 usually double

on segments I-V, single on VI. Hair 7-I,I] well developed, stronger than hair 10

on corresponding segment. Hair 12-I absent. Segment VII: Hairs 1 and 3 usu-

ally triple (2,3b). Comb scales 10-12 (5-16) in single or irregular double row.

Siphon: Index 1.9-2.4; concave on basal ventral surface, inflated at middle, sharp-

ly reduced in diameter apically. Pecten teeth 10-12 (8-15); restricted to basal 0.3

of siphon. Hair 1-S usually triple (3-5b), 0.30-0.35 from base of siphon. Anal

Segment: Hair 1-X usually double (2-4b). Gills usually about 4.5 (3.0-5.0) of sad-

dle length.

SYSTEMATICS. Adults of varipalpus can be distinguished from other species

of the group by the cluster of light setae arising from the subspiracular scale patch

and the absence of postcoxal scales. The presence of setae in the subspiracular

area is an unusual character, known in no other mosquito except laguna and Ae-

des muelleri. The golden scale patch on the anterior mesonotum is relatively sparse,

with golden scales blending gradually into dark brown scales laterally. Lower mep

Arnell & Nielsen: Aedes varipalpus Group 19

setae are usually present. The male genitalia are distinguishable from those of

monticola, laguna and deserticola by the basal tergomesal lobe of the sidepiece

having numerous long, slender bristles in addition to the single differentiated seta

while the former 3 species have 3 or less slender bristles. The pupa of varipal-

pus has a pigmentation identical to that of monticola but it can be distinguished

from the latter by hair 1-IV being more strongly developed than 1-III or 1-V.

The larva of varipalpus can be readily separated from laguna and deserticola by

the characters given in the key. However the larvae of varipalpus and monticola

are almost indistinguishable. The characters given in the key are generally good,

but because of considerable variation are not always reliable. The siphon of vari-

palpus is usually shorter than that of monticola, with an index of 1.9-2.4 in the

former and 2.2-2.5 in the latter. ,

Several individuals of varipalpus were collected in Provo, Utah, which had ab-

normally well-developed larval hairs and extra hairs on the venter of the protho-

rax and mesothorax. These specimens were the only examples of “hairy” larvae

noted in this study, although “hairy” larvae are common in some other groups

of treehole-breeding mosquitoes.

Aedes varipalpus is allopatric from the other species in the group with the ex-

ception of Oak Creek Canyon, Arizona, where its range overlaps with monticola.

Material from this area shows evidence of hybridization between the 2 species.

Hybrids have been found in 2 collections, 1 from a cottonwood treehole at Slide

Rock Campground in which all individuals appear to be hybrids, and the other

from an oak treehole near Manzanita Campground, several miles below the former

site, from which monticola, varipalpus and varipalpus-monticola hybrids were taken.

Oak Creek Canyon is in the upper Gila River drainage and the canyon bottom

is at an elevation of 4200 to 4800 feet. While monticola has been taken at sev-

eral nearby localities at comparable elevations, the nearest record of varipalpus

is at Williams, approximately 40 air miles NW of Oak Creek Canyon and at an

elevation of over 6800 feet. Since Oak Creek Canyon is the only record for vari-

palpus in Arizona outside the Colorado River drainage, it is probable that vari-

palpus has invaded the lower elevations in the Oak Creek Canyon area quite re-

cently. It is surprising that nearly all individuals in which hybridization is evi-

dent show introgression of monticola characters into varipalpus, although monti-

cola is much more abundant and probably better established in the immediate

area. The adults of all but 1 hybrid individual are typical varipalpus in general

appearance but with the monticola characters of a few pcx scales, dark scales

on the ppn and an increased number of dark erect scales on the vertex. The pu-

pae are typical varipalpus and since the larvae of these 2 species are difficult to

distinguish from each other no hybrid characters are evident. One specimen (UCLA

682-24) from the Manzanita Campground collection is typical monticola in gen-

eral appearance but with a reduced number of pcx scales and several ssp setae.

The pupa shows the monticola characters of hairs 1-III-V being subequal. If the

varipalpus-like individuals are F, hybrids, which would seem likely due to their

abundance, then the monticola-like hybrid may be a backcross to monticola. Since

varipalpus characters predominate in supposed F, hybrids, in backcrosses into vari-

palpus, monticola characters may become entirely submerged.

Aedes varipalpus probably invaded the Great Basin from the Colorado River

drainage in southwestern Utah and moved north along the Sevier River and the

margins of Pleistocene Lake Bonneville. It is now found only in isolated popu-

lations in a number of canyons containing cottonwood groves in central and south-

ern Utah.

20 Contrib. Amer. Ent. Inst., vol. 8, no. 2, 1972

BIONOMICS. Females of varipalpus attack humans readily although they have

never been encountered in large numbers. They have been taken biting in Zion

National Park, Utah, at dusk and at midday in the shade. Larvae have been col-

lected from treeholes in scrub oak (Quercus gambelli) and cottonwood (Populus

fremontii) in Arizona and cottonwood, narrow-leaf cottonwood (P. angustifolia),

willow (Salix sp.) and maple (Acer spp.) in Utah. Larvae of varipalpus have been

taken in association with Orthopodomyia signifera near Boulder, Garfield Co.,

Utah, and Oak Creek Canyon, Arizona; with Aedes muelleri at Williams, Hualpai

Mountains and Oak Creek Canyon in Arizona; and with monticola in Oak Creek

Canyon. Aedes varipalpus overwinters in the larval stage in the lower elevations

of southern Utah and Arizona and in the egg stage in the higher elevations of

Arizona and Utah where freezing temperatures are common for several months

of the year.

DISTRIBUTION (fig. 1). Aedes varipalpus has been collected in the Colorado

River drainage of southwestern Utah and northwestern Arizona, at 1 locality in

the Gila River drainage of Arizona and the eastern margins of the Great Basin

in Utah. Material examined: 1178 specimens; 200 males, 484 females, 303 lar-

vae, 191 pupae; 150 individual rearings (134 larval, 11 pupal, 5 incomplete).

UNITED STATES

Arizona. Coconino Co,: Oak Creek Canyon, Manzanita Campground, 4700 ft, 24 May 1970,

HA, LTN, RN (UCLA 682), 2 IpM (682-18,27), 2 IpF (682-16,19), 9 M, 6 F, 2 L [UCLA-

UTAH]. Williams, 6800 ft, 4 Aug 1956, J.N. Belkin and W.A. McDonald (UCLA 212), 9 IpM (212-

143,160,162,165,168,174,186,199,210), 9 IpF (212-161,164,166,170,178,179,197,217,218), 5

pM (212-148,151,153-155), 2 pF (212-145,149), 5 L [UCLA]; same data (UCLA 213), 2 F

[UCLA] ; same data (UCLA 214), 2 IpM (214-116,121), 10 lpF (214-105,111-114,117-120,122),

3 pM (214-107,109,110), 1 pF (214-106), 1 lp (214-115), 10 L [UCLA]; same data, 5 Aug

1956 (UCLA 216), 1 F [UCLA]. Mojave Co.: Beaver Dam, 1 Jan 1967, TIZ (UCLA 416), 5 IpM

(416-10,14,18,20,22), 8 IpF (416-11-13,15-17,19,21), 10 M, 11 F, 23 P, 22 L [UCLA]. Hualpai

Mtn. Park, 6200 ft, 4 Aug 1956, J.N. Belkin and W.A. McDonald (UCLA 210), 3 IpF (210-102,

103,105), 2 Ip (210-101,104), 3 L [UCLA]; same data, 3 Mar 1969, HA, LTN (UCLA 667),

1 IpM (667-16), 9 IpF (667-11-15,17-20), 8 M, 77 F, 35 L [UCLA-UTAH].

Utah. Beaver Co.: Beaver (2 mi E), 5970 ft, 24 Oct 1969, HA (UCLA 675), 6 M, 7 F [UCLA-

UTAH]. Garfield Co.: Boulder (10 mi E), 5600 ft, 17 May 1969, LTN (UCLA 650), 5M,3F

[UCLA-UTAH]. Escalante River at Hwy. 12, 5000 ft, 16 May 1969, LTN (UCLA 649), 9 M,

20 F [UCLA-UTAH]. Jron Co.: Parowan (2 mi S), 6000 ft, 24 Oct 1971, HA (UCLA 676), 7 M,

13 F [UCLA-UTAH]. Kane Co.: Cottonwood Canyon, 22 mi N Hwy. 89, 5000 ft, 26 May

1970 (UCLA 683), 30 M, 49 F, 3 L [UCLA-UTAH]. Orderville, 5460 ft, 25 Mar 1969, HA,

LTN (UCLA 677), 3 IpM (677-13,15,18), 5 lpF (677-11,14,16,17,19), 1 M, 1 F [UCLA-UTAH].

Millard Co.: Oak City (4 mi E), 5900 ft, 20 Aug 1969, LTN (UCLA 651), 3 IpM (651-12,16,18),

3 IpF (651-11,15,17), 1 lp (651-20), 3 M, 15 F [UCLA-UTAH]. Piute Co.: Junction (2 mi W),

7000 ft, 23 Aug 1969, LTN (UCLA 652), 1 IpM (652-16), 5 IpF (652-11,12,15,17,18), 1 Ip

(652-19), 5 M, 24 F [UCLA- UTAH]. Marysvale (1 mi W), 5900 ft, 26 May 1970, HA, LIN,

RN (UCLA 684), 3 IpM (684-12,17,20), 4 IpF (684-11,13,15,19) [UCLA-UTAH] . Utah Co.:

Provo (4 mi N), 4680 ft, 4 Oct 1970, HA (UCLA 698), 2 IpM (698-15-18), 6 IpF (698-12-14,

16,17,19), 6 M, 16 F [UCLA-UTAH]. Washington Co.: Beaver Dam Wash, 2 mi N Motoqua,

4350 ft, 8 Nov 1967, HA, LTN (UCLA 656), 2 M, 3 F [UCLA-UTAH]. Virgin (4.6 mi N),

4000 ft, 25 Mar 1969, HA, LTN (UCLA 673), 2 IpM (673-14,18), 8 IpF (673-11-13,15-17,19,

20), 27 M, 69 F, 12 P, 45 L [UCLA-UTAH]. Leeds, 2750 ft, 25 Mar 1969, HA, LTN (UCLA

674), 2 IpM (674-15,17), 8 IpF (674-11-14,16,18-20), 22 M, 59 F, 24 L [UCLA-UTAH]. Pine

Valley (3 mi E), 6000 ft, 3 June 1970, RN (UCLA 691), 3 IpF (691-13,16,18), 2M,9 F,6P,9L

[UCLA-UTAH]. Zion Natl. Park (Springdale entrance), 3950 ft, 31 Dec 1966, TJZ (UCLA 415),

2 IpM (415-10,11), 2 IpF (415-12,13), 4 L [UCLA]. Zion Natl. Park (Visitor’s Center), 3950 ft,

25 Mar 1969, HA, LTN (UCLA 678), 1 lpM (678-13), 4 lpF (678-11,12,14,18) [UCLA-UTAH] .

Arnell & Nielsen: Aedes varipalpus Group 21

HYBRIDS. Hybrids between varipalpus and monticola are known only from Oak

Creek Canyon, Coconino County, Arizona. For discussion see under systematics

above. Material examined: 65 specimens; 4 males, 22 females, 23 larvae, 16 pu-

pae; 11 individual larval rearings.

UNITED STATES

Arizona. Coconino Co.: Oak Creek Canyon, Slide Rock Campground, 4800 ft, 24 May 1970,

HA, LTN, RN (UCLA 681), 2 IpM (681-14,18), 4 IpF (681-11,13,15,17), 4 L [UCLA-UTAH].

Oak Creek Canyon, Manzanita Campground, 4700 ft, 24 May 1970, HA, LTN, RN (UCLA

682), 1 IpM (682-21), 4 IpF (682-13,17,24,25), 1 M, 14 F, 5 P, 8 L [UCLA-UTAH].

4. Aedes (Ochlerotatus) deserticola Zavortink

Figs. 1,2,10,11

1970. Aedes (Ochlerotatus) deserticola Zavortink, 1970:1-7. TYPE: Holotype male (UCLA

516-32) with associated larval and pupal skins, Littlerock (5 mi SW), San Gabriel Moun-

tains, Los Angeles Co., California, U.S.A., 4 Feb 1969, T.J. Zavortink [USNM].

FEMALE (fig. 2). Wing: 3.75 mm. Proboscis: 2.20 mm. Forefemur: 1.82 mm.

Abdomen: about 2.90 mm. Head: Erect scales yellow on vertex; numerous dark

erect scales laterally. Proboscis entirely dark scaled. Palpus with a few white scales

on apex of segment 1. Thorax: Mesonotum with broad patch of narrow golden

scales from anterior promontory to near prescutellar space, scale patch relatively

sparse, especially just laterad of acrostichal line; outer posterior dorsocentral line

of white and yellow scales with golden scales anteriorly; patches between outer

posterior dorsocentral line and supraalar line golden and yellow posteriorly. Acros-

tichal bristles dark. White scale patches on pleuron relatively small, setae reduced;

ppn entirely white scaled, scales more numerous posteroventrally, 4-8 mixed light

and dark setae posteriorly; ppl with about 10-15 light setae; ssp without setae;

pcx without scales; lower mep with small, compact scale patch in center of scle-

rite; metameron without scales. Hindtarsal segment 1 with broad basal and apical

white rings, white scales occupying about 0.4 of segment; hindtarsal segment 4

with occasional white scales basally and apically. Wing with white scales at base

of C reduced.

MALE. Wing: 2.95 mm. Proboscis: 2.30 mm. Forefemur: 1.70 mm. Abdomen:

about 2.75 mm. Proboscis with ventral white streak on basal 0.5, usually extend-

ing dorsally to form white ring from basal 0.3-0.5. Palpus without white scales

Over joint between segments 2 and 3.

MALE GENITALIA (fig. 10). Segment IX: Tergite lobe with 4-6 (2-7) setae.

Sidepiece: Length 4.0 of width at middle; lateral 0.33 of dorsal surface with strong

bristles and scales, mesal 0.67 of dorsal surface with short bristles, becoming strong-

er toward apex; tergomesal margin bare; basal tergomesal lobe with single strongly

differentiated, apically curved seta and 1-3 long, slender bristles; length of dif-

ferentiated seta about 30.0 of width at base. Claspette: Stem moderately curved,

length 8.0 of width; filament slightly longer than stem, flattened beyond mid-

dle, sharply recurved at apex; connection between claspettes declivous mesally.

Clasper: Strongly curved apically; spiniform about 0.3 of clasper length.

PUPA (fig. 10). Abdomen: 3.37 mm. Trumpet: 0.43 mm. Paddle: 0.79 mm.

Cephalothorax: Light yellowish-brown; trumpet light amber; metanotum light yel-

lowish-brown. Abdomen: Light yellowish-brown. Hair 1-II usually double; 1-[I-V

22 Contrib. Amer. Ent. Inst., vol. 8, no. 2, 1972

subequal. Hair 5-III subequal to 1-III and weaker than 3-III. Hairs 5-I[V-VI strong-

ly developed, subequal. Hair 7-V usually double. Paddle: Apex emarginate; length

1.4-1.6 of width.

LARVA (fig. 11). Head: 0.76 mm. Siphon: 0.70 mm. Saddle: 0.22 mm. Tho-

rax: Integument unpigmented, living larva white. Hair 7-P usually double (2,3b).

Abdomen: Integument as on thorax. Hair 1-II stronger than 13-I]; hairs 1,13-

IlI-V subequal, usually double, with lateral branch longer than medial branch.

Hair 6 usually double on segments I-V, single on VI. Hairs 7-I,1] moderately de-

veloped, subequal to or weaker than hair 10 on corresponding segment. Hair 12-

I present. Segment VIII: Hair 1 usually single (1-3b); hair 3 usually double (1,

2b). Comb scales 7-12 (5-15), in single or irregular double row. Siphon: Index

2.2-2.4 (2.1-2.5); slightly inflated, broadest at level of hair 1-S, sharply reduced

in diameter apically. Pecten teeth 7-11 (4-13); restricted to basal 0.2 of siphon.

Hair 1-S triple, 0.25-0.30 from base of siphon. Anal Segment: Hair 1-X usually

single (1,2b). Gills 4.0-5.0 (3.8-5.5) of saddle length.

SYSTEMATICS. Adults of deserticola differ markedly from the other members

of the group in the reduction of the size of the scale patches on the pleuron,

the scales being more appressed, and in the reduction in the number of pleural

setae. The male genitalia are very similar to laguna and monticola with 3 or few-

er stout bristles in addition to 1 differentiated seta on the basal tergomesal lobe

of the sidepiece, but can be distinguished by the relatively greater stoutness of

the differentiated seta, the length being about 25-30 times the basal diameter,

and the relatively few (4-6) setae on the lobe of the ninth tergite. The pupa of

deserticola is entirely light yellowish-brown without darker areas dorsally as in

varipalpus and monticola. Hair 3-III is generally stronger than 1-III or 5-HI. The

larva of deserticola is unpigmented and hair 12-I is present. It can also be dis-

tinguished by single hairs 1-VIII and 1-X and fewer comb scales (7-11) than in

the other species.

BIONOMICS. Little is known of the habits of deserticola. The larvae have been

collected in treeholes in sycamores (Platanus racemosa), cottonwoods (Populus fre-

montii), live oak (Quercus engelmannii, Q. wislizenii) and scrub oak (Q. turbinella),

and in association with sierrensis and Orthopodomyia signifera. In the Little San

Bernardino Mountains, deserticola has been taken from scrub oak holes less than

1 cm in diameter and holding less than 20 ml of water. In 2 instances larvae

were removed from rotholes to which the only opening was the gallery of a beetle

larva. When larvae of deserticola and sierrensis were reared together in the labor-

atory, the rate of development of deserticola was much slower, but it is not known

whether deserticola is competitively inferior to sierrensis or has an intrinsically

slower growth rate. Nothing is known of the habits of the adults.

DISTRIBUTION (fig. 1). Aedes deserticola is known from the western portions

of the Mojave and Colorado Deserts, the interior slopes of the Transverse and Pen-

insular ranges and the Little San Bernardino Mountains. Material examined: 1703

specimens; 216 males, 203 females, 951 larvae, 333 pupae; 169 individual rear-

ings (155 larval, 14 incomplete).

UNITED STATES

California. Los Angeles Co.: Mt. Emma Rd., 1 mi E Angeles Forest Hwy., 21 Mar 1969, HA,

TJZ, LTN (HA-2-69), 1 IpM (4), 4 M, 17 F [UTAH]. Littlerock (5 mi SW), 4 Apr 1964, TJZ

(UCLA 503), 3 M, 5 F, 144 L [UCLA]; same data, 4 Feb 1969 (UCLA 516), 2 lpM (516-10,34),

17 IpF (516-35-51), 1 lp (516-31), 17 M, 38 P, 85 L [UCLA]; same data (F; gen. from lab

colony) (UCLA 516A), 4 IpM (516A-11-14), 7 IpF (516A-10,16-20,22), 2 Ip (516A-21,23), 51M,

Arnell & Nielsen: Aedes varipalpus Group 23

53 F [UCLA]; same data (F2 gen. from lab colony) (UCLA 516B), 8 lpM (516B-10,11,13-18),

2 Ip (516B-12,19), 240 L [UCLA]; same data (UCLA 517), 4 IpM (517-30,31,33,34), 17 IpF

(517-35-39,41-43,45-53), 1 lp (517-40), 18 M, 14 F, 39 P, 102 L [UCLA]; same data (UCLA

522), 4 lpM (522-10-13), 4 lpF (522-15-17,19), 11 M, 4 F, 13 P, 8 L [UCLA]. Pearblossom (8

mi SE), 4 Feb 1969, TJZ (UCLA 518), 2 IpM (518-21,25), 3 IpF (518-28-30), 8 L [UCLA];

same data, 12 Mar 1969, TJZ, DWH (UCLA 530), 1 IpM (530-12), 4 IpF (530-10,11,14,15), 1 M,

1 F, 2 P, 1 L [UCLA]. Riverside Co.: Anza (11 mi E, Santa Rosa Mts.), 6700 ft, 20 July 1970,

TJZ (UCLA 574), 1 lpM (574-21) [UCLA]. Joshua Tree Natl. Mon., Juniper Flats, 4900-5100

ft, 17 Mar 1969, TJZ, DWH (UCLA 539), 18 IpM (539-10-27), 5 IpF (539-28-32), 15 M, 12 F,

32 P, 119 L [UCLA]; same data (UCLA 545), 4 IpM (545-10,11,16,17), 3 IpF (545-14,18,19),

3 Ip (545-12,13,15), 2 M, 4 F, 6 P, 4 L [UCLA]; same data (UCLA 546), 2 IpM (546-10,12),

1 IpF (546-14), 3 Ip (546-11,13,15) [UCLA]. Joshua Tree Natl. Mon., Hidden Valley, 20 May

1946, A.L. Melander, 1 F [USNM]. San Bernardino Co.: Yucca Valley (6.5 mi NW), 4400 ft,

18 Mar 1969, TJZ, DWH (UCLA 544), 3 lpM (544-10,11,13), 2 IpF (544-14,15), 1 lp (544-12)

[UCLA] ; same data (UCLA 548), 1 lpM (548-13), 4 IpF (548-10-12,14) [UCLA]. Yucca Valley

(9 mi NW), 4600 ft, 18 Mar 1969, TJZ, DWH (UCLA 547), 1 L [UCLA]. San Diego Co.:

Banner Canyon (5 mi E Julian), 12 Feb 1969, TJZ (UCLA 527), 13 lpM (527-23-25,27-32,34,

37-39), 15 IpF (527-22,26,33,35,36,40-49), 24 M, 7 F, 34 P, 71 L [UCLA]; same data (UCLA

528), 2 IpM (528-20,21), 3 IpF (528-22,24,25), 1 lp (528-23) [UCLA].

5. Aedes (Ochlerotatus) sierrensis (Ludlow)

Figs13242,13

1905. Taeniorhynchus sierrensis Ludlow, 1905:231-232. TYPE: Lectotype female, Three Rivers,

Sierra Nevada (Tulare Co.), California, U.S.A., E.J. Bingham [USNM; designation by

Stone and Knight, 1956:225].

Aedes (Ochlerotatus) sierrensis of Belkin and McDonald (1957:191); Myers (1967:805); Zavortink

(1970:1); Lunt and Nielsen (1971:110).

Aedes varipalpus of Dyar (1907:125; 1917a:18; 1917b:99; 1919:39; 1920a:19); Howard, Dyar

and Knab (1913:plates 31,75); Matheson (1929:139,233); Reeves (1940:39); Stage, Gjullin

and Yates (1952:60,74).

Aedes sierrensis of Nielsen, Linam and Rees (1963:426); Nielsen, Arnell and Linam (1967:75);

Nielsen, Linam, Arnell and Zavortink (1968:362).

Aedes (Finlaya) sierrensis of Rohlf (1963:802); Trebatoski and Haynes (1969:332); Cupp and

Horsfall (1969:51).

Aedes varipalpus in part of Howard, Dyar and Knab (1917:644); Jenkins and Carpenter (1946:

39); Peyton (1956:220); Richards, Nielsen and Rees (1956:14).

Aedes (Finlaya) varipalpus of Matheson (1944:188,291).

Aedes (Finlaya) varipalpus in part of Edwards (1932:154); Carpenter and LaCasse (1955:257).

Aedes (Taeniorhynchus) varipalpus of Dyar (1918:74; 1920b:171; 1928:214,524); Hearle (1926:

53). :

Aedes (Taeniorhynchus) varipalpus in part of Dyar (1918:74; 1922:86).

Aedes (Culicelsa) varipalpus of Freeborn (1926:416).

Grabhamia varipalpus of Dyar (1905a:48; 1905b:53).

Culex varipalpus of Dyar (1904a:39; 1904b:90); Blaisdell (1906:107).

Culicada varipalpus of Felt (1905:447).

Taeniorhynchus (?) sierronsis of Theobald (1910:432).

FEMALE (fig. 2). Wing: 3.50 mm. Proboscis: 2.30 mm. Forefemur: 1.75 mm.

Abdomen: about 2.85 mm. Head: Erect scales yellow on vertex, dark laterally.

Proboscis entirely dark scaled or occasionally with scattered white scales ventral-

ly. Palpus often with a few white scales on apex of segment 1. Thorax: Meso-

24 Contrib. Amer. Ent. Inst., vol. 8, no. 2, 1972

notum with broad, dense patch of narrow golden scales from anterior promon-

tory to near prescutellar space or with narrow line of golden scales along acros-

tichal line from anterior promontory to near prescutellar space, golden scales be-

ing reduced in number laterad to acrostichal line but forming indistinct line just

mesad to anterior dorsocentral line; lateral prescutal white scales relatively sparse;

outer posterior dorsocentral line broad anteriorly, composed mostly of white scales

with a few golden scales, narrowed posteriorly and composed of yellow scales;

patches between outer posterior dorsocentral line and supraalar line relatively in-

conspicuous, golden anteriorly, white or yellow posteriorly. Acrostichal bristles

dark. Pleural setae reduced; ppn with narrow dark scales anteriorly and broader

white scales in sparse patch posteriorly, scales becoming more dense posteroven-

trally, 5-8 mixed light and dark setae posteriorly; ppl with 10-15 light setae; ssp

without setae; pcx without scales; lower mep scale patch compact, scales not reach-

ing ventral 0.33 of sclerite. Hindtarsal segment 1 with broad basal and apical white

rings, white scales occupying 0.4 of segment; hindtarsal segment 4 with broad

ring of white scales on basal 0.3 of segment and a few white scales apically. Wing

with white scales on base of C reduced.

MALE. Wing: 2.80 mm. Proboscis: 2.25 mm. Forefemur: 1.65 mm. Abdomen:

about 2.55 mm. Proboscis entirely dark scaled or often, in specimens with white

scales over joint between palpal segments 2 and 3, with scattered white scales

ventrally from basal 0.25-0.60, these rarely forming indistinct ring. Palpus with

or without white scales over joint between segments 2 and 3.

MALE GENITALIA (fig. 12). Segment [X: Tergite lobe with 4-7 (2-9) setae.

Sidepiece: Length 3.0 of width at middle; basal 0.75 of dorsal surface covered

with short bristles, apical 0.25 with strong bristles; tergomesal margin with num-

erous short, stout bristles; basal tergomesal lobe with dense patch of strongly de-

veloped, apically curved setae. Claspette: Stem moderately curved, length 4.0 of

width; filament slightly longer than stem, bluntly recurved at apex; connection

between claspettes nearly horizontal but with median emargination. Clasper: Slight-

ly curved apically; spiniform 0.25 of clasper length.

PUPA (fig. 12). Abdomen: 3.25 mm. Trumpet: 0.44 mm. Paddle: 0.83 mm.

Cephalothorax: Light brown, darker dorsally; trumpet amber; metanotum light

brown, haltere cases lighter. Abdomen: Light brown, anterior segments darker,

especially mesally. Hair 1-II usually double; 1-IV usually slightly stronger than

1-III and 1-V. Hair 5-III subequal to 1-IIJ and weaker than 3-III. Hair 5-VI weak-

er than 5-IV,V which are subequal. Hair 7-V usually double. Paddle: Apex emar-

ginate, occasionally rounded; length 1.6-1.8 of width.

LARVA (fig. 13). Head: 0.82 mm. Siphon: 0.83 mm. Saddle: 0.28 mm. Tho-

rax: Integument pigmented, living larva gray in color. Hair 7-P usually double,

rarely triple. Abdomen: Integument as on thorax. Hair 1 weaker and with more

branches than hair 13 on segments IIJ-V. Hair 6 double (2,3b) on segments I-IV,

double on V and single on VI. Hairs 7-I,II moderately developed, subequal to

or weaker than hair 10 on corresponding segment. Hair 12-I present. Segment

VI: Hair 1 usually 4b (2-5); hair 3 usually 4b (3-5). Comb scales 14-20 (7-24),

in irregular double or triple row. Siphon: Index 2.7-3.3; sides nearly parallel, not

sharply reduced in diameter apically. Pecten teeth 12-14 (8-19); restricted to basal

0.30 (0.23-0.30) of siphon. Hair 1-S usually triple (3,4b), 0.35-0.40 from base

of siphon. Anal Segment: Hair 1-X usually double (1-3b). Gills 3.0-4.0 of saddle

length.

SYSTEMATICS. Adults of sierrensis differ from the remaining members of the

Arnell & Nielsen: Aedes varipalpus Group 25

group primarily in the reduction of pleural setae, the broad basal white ring on

hindtarsal segment 4, the absence of the ventral white streak on the male pro-

boscis, and the male genitalia. The sidepiece of the male genitalia is relatively

short and stout; the basal 0.75 of the dorsal surface is covered with short hairs

(the remaining members of the group have long hairs and scales on the lateral

0.33 of the dorsal surface); the tergomesal margin has numerous short, strong

hairs (bare in the remaining species); and the basal tergomesal lobe is charact-

erized by a dense patch of strongly developed setae (in contrast to the single

strong seta of the other members of the group). The clasper of sierrensis is rel-

atively short and not strongly curved apically and the claspette is relatively short

and thick. The pupa of sierrensis differs from the remaining members of the group

in the shape of the paddle; the dark pigmentation of the abdomen on the first

4 or 5 segments; hair 3-III stronger than 1-III or 5-III, 1-IV stronger than 1-III,V,

and 5-VI weaker than 5-IV,V. The larva of sierrensis is pigmented and light gray

in color. Hair 12-I is present. It shows a slight reduction in the development of

some setae, primarily hairs 1,13-I-V and 7-I,II, an increase in the number of comb

scales (usually 14-20) and a relatively long, narrow siphon.

There is considerable variation in the ornamentation of the adults of sierrensis.

Southern populations are characterized by the presence of a white patch of scales

over the joint between segments 2 and 3 of the male palpus. All material ex-

amined from the Transverse and Peninsular Mountain Ranges of southern Califor-

nia exhibit this character and in collections from the South Coast Range of San

Luis Obispo and southern Monterey Counties some males lack the patch and others

have a small one. All material from northern Monterey County, the San Joaquin

Valley and the Sierra Nevada (including the Tehachapi Range) is without the white

scale patch. The density of the. golden scales on the anterior mesonotum is an

extremely variable character in sierrensis, ranging from a uniformly very dense

condition to a reduction in golden scales just laterad of the acrostichal line which

results in 3 golden stripes rather than a broad golden patch. The latter condition

approaches that found in laguna. A reduction in golden scales is common in sier-

rensis from the eastern Transverse Ranges and Peninsular Ranges of California,

and is found, although to a limited extent, in specimens from throughout the

entire range of the species.

Much of the present distribution of sierrensis has occurred in relatively recent

geological time. Most of the area of the present northern and eastern range of

sierrensis was glaciated during the most recent Pleistocene glacial advance when

the Cordilleran Ice Sheet pushed southward from Canada covering much of the

northern Rocky Mountains west of the Continental Divide, much of the Colum-

bia Plateau and the coastal mountain ranges of British Columbia, Washington and

Oregon. This area must then have been invaded by sierrensis since the Pleisto-

cene, probably within the last 15,000 to 20,000 years. The northern Utah popu-

lation of sierrensis is located in the Great Basin with no drainage connection

to the Columbia system. It is probable that sierrensis invaded the Great Basin

at the maximum extent of Lake Bonneville, a pluvial lake covering the eastern

Great Basin, when there was an outlet to the Snake River in southeastern Idaho.

BIONOMICS. Females of sierrensis are avid biters of humans in central and

northern California, constituting a major pest species in localized areas and the

object of control programs. However there is some evidence that populations of

sierrensis in Utah, southern California and British Columbia are not readily at-

tracted to humans. Nothing is known of the natural hosts for this species other

26 Contrib. Amer. Ent. Inst., vol. 8, no. 2, 1972

than man, although in the laboratory it will readily feed on rabbits and chickens.

Aedes sierrensis has been shown to be capable of transmitting Western Equine

Encephalitis under laboratory conditions (Stage, Gjullin and Yates, 1952). Mating

usually occurs near the host. Males of sierrensis are attracted to the host animal

where they await the arrival of females and mating takes place in the air or while

the females are taking a blood meal.

Larvae of sierrensis are found exclusively in treeholes or situations resembling

treeholes. They seem to have no preference for any particular species of tree and

are found wherever suitable treeholes form. In the extensive forests and wood-

lands of the Pacific Coast, sierrensis is found in a wide variety of trees, but in

the eastern, more arid regions of its range, it is found exclusively in riverine sit-

uations in cottonwood (Populus). Aedes sierrensis has been collected in associ-

ation with deserticola and Orthopodomyia signifera in California and with Aedes

hendersoni Cockrell, 1918 in British Columbia, Montana and eastern Oregon.

Eggs are laid singly on the sides of treeholes above the water line. Hatching

occurs when water in the treeholes rises to flood the eggs, although there is evi-

dence that eggs may drop off the sides of the hole into the water. As in the

case with the other members of the Varipalpus Group and treehole aedines in

general, not all eggs hatch the first time they are flooded, some hatching only

after repeated wetting and drying. Judson, Hokama and Kliewer (1966) in studies

on the embryogeny and hatching of sierrensis eggs determined that 9-15 days are

required for embryonation and that the egg had an uncommonly thick chorion

and serosal cuticle. A strong hatching stimulus, an unusually low level of dissolved

oxygen, was shown to be necessary for eclosion. Eggs will hatch at very low tem-

peratures as Arnell has observed first instar larvae in treeholes nearly filled with

ice. The rate of larval development varies with temperature with complete devel-

opment from egg to adult varying from as little as 14 days or as long as several

months. Larvae hatched by early fall rains in California will often not pupate in

significant numbers until early spring. This period of dormancy is apparently re-

lated to photoperiod as changes in temperature under laboratory conditions fail

to have any effect. Cold temperatures apparently retard pupation. In Utah, no

pupae of sierrensis have been found later than October when water temperatures

drop below approximately 50° F, even though larvae have been found through-

out the winter in treeholes covered with several inches of ice and with water tem-

peratures as low as 30° F. Pupation begins almost immediately after these larvae

are brought into the laboratory and the water temperature is raised. In the milder

climates of the Pacific Coast larvae are unable to survive prolonged periods of

freezing temperatures.

DISTRIBUTION (fig. 1). Aedes sierrensis has the most extensive range of all

the members of the group. It has been collected throughout the Pacific Coast

drainage systems from southern California to British Columbia including Montana

and Idaho, the desert margins of southern California, the eastern slope of the

Sierra Nevada in California and Nevada and the Great Basin in northern Utah.

Material examined: 10,504 specimens; 1904 males, 1433 females, 5014 larvae,

2153 pupae; 637 individual rearings (505 larval, 107 pupal, 25 incomplete).

CANADA

British Columbia. Caulfeild, 19 July 1919, Eric Hearle, 3 F [USNM]. Harrison, 20 June

1919, Eric Hearle, 1 M, 1 F [USNM]. Karemos (4 mi W), 4 Aug 1968, LTN (UCLA 647), 3 M,

4 F, 6 L [UCLA-UTAH]. Kaslo, 13 June 1903, H.G. Dyar, 17M, 14 F,1P,8L,E [USNM] ;

same data, Apr 1904, 5 L [USNM]. Kelowna, 3 Aug 1968, LTN (UCLA 646), 2 M,4F, 2 L

Arnell & Nielsen: Aedes varipalpus Group 2a

(UCLA: UTAH]. Kootenay Lake (S end), 1 Aug 1968, LTN (UCLA 645), 7 F, 2 L [UCLA-

UTAH]. Mara Lake, 2 Aug 1968, LTN (UCLA 648), 2 M,5 F, 1 L [UCLA-UTAH]. Mission,

14 July 1920, H.G. Dyar, 1 F [USNM]. Nanaimo, 6 Aug 1906, Dyar and Caudell, 2 M, 2 F

[USNM]. Terrace, 13-14 Aug 1919, H.G. Dyar, 6 M, 28 F [USNM]. Vancouver (Stanley Park),

6 Aug 1906, Dyar and Caudell, 1 M [USNM]. Vancouver Island, 8 Aug, K. Taylor, 5 M, 10 F

[USNM]. Victoria, 12 Aug 1906, Dyar and Caudell, 3 F sniieiadl Wellington, 8 Aug 1906,

Dyar and Caudell, 2 F [USNM].

‘UNITED STATES

California. Alameda Co.: Berkeley, 16 Mar 1950, R.E. Ryckman, 3 F [USNM]. Oakland,

20 Mar 1940, H.F. Gray, 5 L [UCLA]. Sunol, Dec 1947, W.W. Wirth, 2 P, 57 L [USNM]; same

data, 4 Feb 1948, 34 L [USNM] ; same data, 24 Mar 1949, 4 M [USNM] . Alpine Co.: Markleeville,

5526 ft, 22 June 1970, LTN, RN, S.J. Carpenter (HA-17-70), 1 M, 3 F, 4 P, 6 L [UTAH].

El Dorado Co.: El Dorado Hills (2.5 mi E), 1000 ft, 5 May 1970, HA, RN (UCLA 680), 3 lpM

(680-16,19,20), 6 IpF (680-12,15,17,18) [UCLA-UTAH]. Kyburz (3 mi W), 4950 ft, 20 June

1970, LIN, RN (UCLA 692), 4 lpF (692-14,15,17,20), 1 Ip (692-19), 2 M, 6 F, 7 L [UCLA-

UTAH]. Placerville, 1900 ft, 21 June 1970, LTN, RN (UCLA 693), 1 IpM (693-19), 4 IpF

(693-13,14,16,18), 6 M, 3 F, 6 L [UCLA-UTAH]. Fresno Co.: Fresno, 13 Apr 1953, 1M

[USNM]. Kearney Park, 22 Apr 1954, 1 M [USNM]. Kings Canyon, 9 July 1955, R.X. Schick,

1 F [UCLA]. Humboldt Co.: Bair’s Ranch, Redwood Cr., 11 June 1903, H.S. Barber, 2 M, 1 F

[USNM]. Eureka, 3 June 1903, H.S. Barber, 1 M [USNM]. Fieldbrook, 30 May 1903, HLS.

Barber, 7 M, 2 F [USNM]. Orleans, Dillon Creek, 3 Sept 1933, CCC Survey, 8 F [USNM].

Prairie Creek, 10 Aug 1948, W.W. Wirth, 1 M [USNM]. Richardson, 23 July 1954, A.L. Melander,

6 M, 2 F [USNM]. Willow Creek (3-4 mi W), 9 Sept 1950, J.N. Belkin (UCLA 72), 3 F [UCLA].

Kern Co.: Bakersfield, 12 May 1948, D. Brockman, 1 F [UCLA]; same data, 26 May 1948, 5 F

' [UCLA]. Bakersfield (6 mi NW), 26 May 1948, W.C. Reeves, 4 F [UCLA]. Tehachapi (7.3 mi

SE), 7 Feb 1969, TJZ (UCLA 525), 5 IpM (525-10-14), 5 lpF (525-15-19), 1 M, 3 P, 3 L

[UCLA]. Locality unknown, 26 May 1948, W.C. Reeves, A. Rudnick, 6 F [UCLA]. Los Angeles

Co.: Acton (6 mi SW), 11 Dec 1965, TJZ (UCLA 508), 2 L [UCLA]. Agoura, 17 May 1952,

McDonald, 1 F [UCLA]. Claremont, Baker, 2 F [USNM]. Gorman (10.6 mi E, 1.8 mi NNW

Hwy. 138), 7 Feb 1969, TJZ (UCLA 523), 1 lpF (523-10), 3 M, 3 P, 2 L [UCLA]. Littlerock

(5 mi SW), 4 Apr 1964, TJZ (UCLA 503), 1 M, 3 F, 42 L [UCLA]; same data, 4 Feb 1969

(UCLA 516), 1 lpM (516-28), 18 IpF (516-11,13-27,29,30), 53 M, 26 F, 87 P, 90 L [UCLA];

same data (UCLA 517), 6 IpM (517-10-15), 14 IpF (517-16-29), 60 M, 99 F, 166 P, 83 L

[UCLA]; same data (UCLA 522), 2 IpF (522-14,18), 4 F, 5 P, 3 L [UCLA]. Los Angeles,

1 Feb 1950, G. Heid, 1 M, 1 F, 1 L [UCLA]. Malibu Canyon, 3 Mar 1955, J.N. Belkin (UCLA

158), 1 IpM (158-112), 4 IpF (158-113,116,118,119), 21 M, 1 F, 3 P, 23 L [UCLA]; same data

25 Mar 1956, W.A. McDonald, 1 L [UCLA]. Mt. Emma Road, 1 mi E Angeles Forest Hwy.,

21 Mar 1969, HA, LTN, TJZ (HA-2-69), 5 lpM (3,5,8,9,12), 8 IpF (1,6,7,10,11,13-15), 2 M, 33 F

[UTAH]. Pasadena, Arroyo Seco, 22 May 1906, Dyar and Caudell, 2 F [USNM]. Pearblossom

(4 mi SE), 3600 ft, 12 Mar 1969, TJZ (UCLA 533), 1 lpM (533-10), 146 L [UCLA] ; same data

(UCLA 532), 18 L [UCLA]. Pearblossom (4.5 mi SE), 3700 ft, 12 Mar 1969, TJZ (UCLA 531),

1 IpM (531-10), 1 IpF (531-11), 34 L [UCLA]. Pearblossom (5 mi SE), 4 Feb 1969, TJZ

(UCLA 518), 1 IpM (518-13), 15 IpF (518-10-12,14-20,22-24,26,27), 58 M, 17 F, 70 P, 81 L

[UCLA]; same data (UCLA 519), 12 lpM (519-10-13,15,17,18,21-23,25,29), 8 IpF (519-14,16,

19,20,24,26-28), 262 M, 51 F, 347 P, 864 L [UCLA]. Pearblossom (6.5 mi SE), 4100 ft, 12 Mar

1969, TJZ (UCLA 530), 1 IpF (530-13), 159 L [UCLA]. San Gabriel River, 29 June 1915,

C.F. Adams, 1 M, 1 F [USNM]. Santa Monica Mts., Fall 1951, 17 P [UCLA]; same data,

11 Mar 1954, 3 L [UCLA]; same data, 2 Mar 1957, J. Pumphrey, 14 M, 10 F [UCLA]; same

data, 12 Apr 1957, Bimantoro, 6 M [UCLA]. Sepulveda Canyon, 3 Mar 1956, W.A. McDonald

(UCLA 182), 4 M, 2 F, 31 P [UCLA]. Soledad Canyon, 20 Mar 1955, W.A. McDonald (UCLA

162), 4 IpM (162-109,125,135,137), 7 IpF (162-100-114,126,128), 18 pM (162-101-105,107,

115-119,121,123,124,130,132-134), 5 pF (162-106,108,120,122,131), 1 M, 12 P, 5 L [UCLA].

Tapia Park, 11 Mar 1954, 5 L [UCLA]; same data, 7 May 1954, 35 P, 7 L [UCLA]; same data,

4 Apr 1954, J.N. Belkin (UCLA 102), 1 lpM (102-104), 13 IpF (102-101,102,105-108,110-116),

28 Contrib. Amer. Ent. Inst., vol. 8, no. 2, 1972

1 lp (102-103), 1 L [UCLA]; same data, 7 Apr 1954, J.N. Belkin and W.A. McDonald, 3 L

[UCLA]; same data, 19 Oct 1963, TJZ (UCLA 417), 4 IpM (417-10,13,15,19), 5 IpF (417-12,

14,16-18), 10 M, 2 F, 9 L [UCLA]; same data, 17 Dec 1963 (UCLA 491), 4 M, 3 F [UCLA];

same data (UCLA 489), 3 M, 3 F [UCLA]; same data, 2 Dec 1967 (UCLA 433), 2 IpM (433-

12,14), 15 lpF (433-11,13,14,16-20,22-24,26,29), 3 Ip (433-10,20,25), 101 M, 34 F, 150 P,

665 L [UCLA]. Westwood Hills, 1 Feb 1950, 13 M, 4 F [UCLA]; same data, 6 May 1955, C.

Blodgett, 1 F [UCLA]. Locality unknown, 22 Mar 1952, 19 L [UCLA]. Marin Co.: Ft. Baker,

6 Apr 1953, Schuessler, 1 M, 4 F [UCLA]. Mill Valley, CCC Survey, 3 F [USNM]. Muir Woods

State Park, 24 Feb 1952, 1 F [UCLA]. Mariposa Co.: El Portal, 12 May 1916, H.G. Dyar, 1 F

[USNM]. Yosemite, 16 May 1916, H.G. Dyar, 71 M, 23 F [USNM]. Yosemite Natl. Park, S

entrance, 4800 ft, 13 Sept 1969, HA, LTN (UCLA 668), 1 M [UCLA]. Yosemite Natl. Park,

Yosemite Valley, 21 July 1954, W.A. McDonald, 1 F [UCLA]. Mendocino Co,: Orr’s Hot Springs,

July 1921, Dorothy Dyar, 1 M [USNM]. Monterey Co.: Hunter-Liggett Mil. Res., W boundary,

20 mi W Lockwood, 1600 ft, 27 Oct 1971, HA (UCLA 590), 5 IpM (590-13-15,24,27), 13 IpF

(590-10-12,16-19,21-23,25,26,29), 2 lp (590-20,28), 11 M, 25 P, 40 L [UCLA]. Toro Regional

Park, 5 mi W Salinas, 50 ft, 27 Oct 1971, HA (UCLA 589), 9 IpM (589-14,16,17,19,21,24,25,

28,29), 11 lpF (589-10-13,15,18,20,22,23,26,27), 1 lp (589-22), 10 M, 1 F, 26 P, 121 L [UCLA].

Napa Co.: Bothe-Napa Valley State Park, 500 ft, 16 Sept 1969, HA, LTN, S.J. Carpenter (UCLA

670), 2 lpM (670-15,16), 8 IpF (670-11-14,17-20), 14 M, 36 F, 12 P, 26 L [UCLA-UTAH].

Orange Co.: Irvine Park, 24 June 1949, 1 M, 3 F [UCLA]. Los Alamitos, 4 July 1949,3 M,6F

[UCLA]. Placer Co.: Auburn, 20 Oct 1918, L. Bruner, 1 F [USNM]. Lake Tahoe, 15 Aug 1915,

H.G. Dyar, 1 M, 2 F [USNM]. Plumas Co.: Chester, July 1957, C.L. Hogue, 1 F [UCLA].

Riverside Co.: Aguanga (7 mi NW), 3 Feb 1966, TJZ (UCLA 292), 6 IpM (292-10-12,15,16,20),

3 IpF (292-14,18,19), 16 pM (292-30,31,100,102-114), 24 M, 5 F, 34 P, 80 L [UCLA]. Anza

(11 mi E, Santa Rosa Mts.), 6700 ft, 20 July 1970, TJZ (UCLA 574), 1 pM (574-20) [UCLA].

Elsinore (11 mi SW), 12 Feb 1964, TJZ (UCLA 501), 1 lpM (501-10), 3 M, 1 F, 2 L [UCLA].

Idyllwild, 30 May 1939, 1 F [UCLA]; same data, 18 June 1952, 1 F [UCLA]. Idyllwild (11.5 mi

N), 5000 ft, 2 Nov 1971, HA (UCLA 592), 8 lpM (592-10,15,17,21-23,28,29), 10 IpM (592-12-14,

16,18-20,24-26), 2 Ip (592-11,27), 42 M, 8 F, 57 P, 123 L [UCLA]. Banning (6 mi S), 4000 ft,

2 Nov 1971, HA (UCLA 591), 8 IpM (591-14,15,18,19,22,23,27,29), 11 IpF (591-10-13,16,17,21,

24-26,28), 1 lp (591-20), 8M, 8 P, 14 L [UCLA]. San Jacinto Mts., 1 July 1952, J.F. Powers, 2 F

[UCLA]. Fern Basin, San Jacinto Mts., 15 June 1940, C.D. Michener, 1 F [UCLA]. Sacramento

Co.: Sacramento, 12 Apr 1948, R.M. Bohart, 1 M [USNM]. San Bernardino Co,: Cajon Jct. (1 mi

E), 4 Feb 1969, TJZ (UCLA 520), 1 lpM (520-16), 9 IpF (520-10-15,17-19), 13 M, 1 F, 10 P, 52 L

[UCLA]. Crestline (4 mi NW), 4 Feb 1969, TJZ (UCLA 521), 2 IpM (521-11,14), 8 IpF (521-10,

12,13,15-19), 37 M, 5 F, 48 P, 113 L [UCLA]. San Diego Co.: Banner Canyon, 5 mi E Julian,

12 Feb 1969, TJZ (UCLA 526), 4 M, 5 P, 5 L [UCLA]; same data (UCLA 527), 5 IpM (527-10,

12-14,17), 6 lpF (527-15,16,18-21), 1 lp (527-11), 76 M, 8 F, 87 P, 505 L [UCLA]; same data

(UCLA 528), 4 IpM (528-10-12,14), 6 IpF (528-13,15-19), 11 M, 2 F, 14 P, 91 L [UCLA] ; same

data (UCLA 529), 8 IpM (529-10-14,17-19), 2 lpF (529-15,16), 29 M, 1 F, 36 P, 136 L [UCLA].

Barrett Jct., 1 Feb 1966, TJZ (UCLA 287), 6 L [UCLA] ; same data (UCLA 288), 3 IpM (288-10,

12,15), 2 lpF (288-13,14), 1 lp (288-11) [UCLA]; same data (UCLA 289), 2 IpM (289-11,15),

9 IpF (289-10,12-14,16-20), 14 pM (289-100-113), 2 L [UCLA]. Bonsall, 12 Feb 1964, TJZ (UCLA

498), 10 M, 4 F [UCLA] ; same data (UCLA 499), 2 F [UCLA] ; same data (UCLA 513), 19 M, 11 F,

14 L [UCLA]. Campo (5.1 mi NE), 1 Jan 1966, TJZ (UCLA 512), 3 L [UCLA]. Campo (4.9 mi

NE), 2 Feb 1966, TJZ (UCLA 290), 9 lpM (290-10-18), 15 pM (290-31-34,100-111,113), 3 pF

(290-30,112,114), 1 lp (290-19), 10 M, 4 F, 18 P, 131 L [UCLA]. Jacumba, 2 Feb 1966, TJZ

(UCLA 291), 5 IpM (291-10,11,13-15), 2 IpF (291-16-18), 13 pM (291-30,100-110,112,113), 3 pF

(291-31,33,111), 4 M, 10 F, 12 P, 24 L [UCLA]. Pala (0.5 mi W), 3 Feb 1966, TJZ (UCLA 293),

2 IpM (293-10,18), 7 IlpF (293-11,13,16,17,19,31), 17 pM (293-40,41,100-114), 2 pF (293-30,42),

32 M, 5 F, 43 P, 79 L [UCLA]. Pauma Valley (18 mi SE), 1 Jan 1966, TJZ (UCLA 510), 3 L

[UCLA]; same data (UCLA 511), 3 L [UCLA]. San Joaquin Co.: Stockton, H.J. Quayle, 1 F

[USNM]. San Luis Obispo Co.: La Panza Campground, 28 mi E Santa Margarita, 2400 ft, 18 Apr

1964, TJZ (UCLA 505), 15 M, 14 F [UCLA]; same data, 26 Oct 1971, HA (UCLA 587), 5 lpM

Arnell & Nielsen: Aedes varipalpus Group 29

(587-13,15,16,20,27), 13 IpF (587-10-12,14,17-19,22-26,28), 2 lp (587-21,29), 13 M, 4 F, 27P,

51 L [UCLA]. La Panza Campground (3 mi E), 2500 ft, 26 Oct 1971, HA (UCLA 586), 3 IpF

(586-10-12) [UCLA]. La Panza Ranger Station, 27 mi E Santa Margarita, 2400 ft, 26 Oct 1971, HA

(UCLA 588), 7 lpM (588-10-13,15-17), 1 lp (688-14) [UCLA] . San Mateo Co.: La Honda, 26 Mar

1949, F. Prince, 15 M, 2 F [UCLA]. Santa Barbara Co.: Santa Barbara (2 mi N), 50 ft, 25 Sept

1971, HA (UCLA 583), 2 IpM (583-13,15), 3 IpF (583-11,12,17), 2 lp (583-10,14) [UCLA] . Sol-

vang (5.5 mi SW), 700 ft, 25 Sept 1971, HA (UCLA 584), 3 IpM (584-18,23,32), 27 IpF (584-

10-17,19-22,24-31,33-39), 11 M, 2 F, 37 P, 129 L [UCLA]. Santa Clara Co.: Palo Alto (Stanford),

23 June-7 July 1903, 4 M, 9 F [USNM]; same data, 27 Dec 1965, TJZ (UCLA 509), 2 L [UCLA].

San Jose, 2 Mar 1941, W.C. Reeves, 8 M [UCLA]. Santa Cruz Co.: Big Basin, 18 July 1954, A.L.

Melander, 3 M [USNM]. Shasta Co.: Hazel Creek, 9-10 Sept 1933, CCC Survey, 14 F [USNM].

McArthur-Burney Falls St. Park, 3000 ft, 18 Sept 1969, HA, LTN (UCLA 671), 4 lpM (671-11,

14,16,19), 6 lpF (671-12,13,15,17,18,20), 12 M, 13 F, 3 L [UCLA-UTAH] . Siskiyou Co.: Dunsmuir,

19 July 1906, Dyar and Caudell, 5 F [USNM]. Sonoma Co.: Monte Rio, 16 Feb 1952, 6 M

[UCLA]. Tehama Co.: Corning, 2 Apr 1957, R. Gifford, 3 F [UCLA] . Tulare Co.: Sequoia Natl.

Park, Heather Lake, 8 Aug 1949, W.W. Wirth, 1 M [USNM]. Three Rivers, 800 ft, 13 Sept 1969,

HA, LTN (UCLA 669), 3 IpM (669-16-18), 7 IpF (669-11-15,19-21), 28 M, 33 F, 12 P, 34 L

[UCLA-UTAH] ; same data, E.J. Bigham, 1 M, 10 F [USNM]. Tuolumne Co.: Yosemite Natl. Park,

Tuolumne Grove, 22. July 1954, W.A. McDonald, 2 F [UCLA]. Ventura Co,: Lake Piru, 9 Feb

1964, TJZ (UCLA 496), 119 M, 39 F, 27 P, 68 L [UCLA] ; same data (UCLA 418), 2 M [UCLA].

Yolo Co.: Davis, 2 June 1952, A.D. Telford, 16 L [USNM] . County Unknown: Nice, 22 Feb 1942,

A.W. Lindquist, 1 M, 1 P, 4 L [USNM]. Taylor Park Estates, 15 Feb 1952, 1 F [UCLA]. Locality

Unknown: Dyar and Caudell, 5 M, 2 F [USNM].

Idaho. Bingham Co.: Firth, 4550 ft, 13 June 1970, HA (UCLA 690), 1 IpM (690-19), 4 IpF

(690-11,13,14,17), 1 Ip (690-12), 17 M, 39 F, 14 L [UCLA-UTAH]. Blaine Co.: Bellevue (2.5 mi

S), 5300 ft, 25 July 1969, JHL (UCLA 672), 2 IpM (672-13,14), 3 IpF (672-11,12,17), 10 M, 2 F

[UCLA-UTAH]. Ketchum (6 mi N), 6200 ft, 28 July 1968, LTN (UCLA 644),4M,5 F,2P,2L

[UCLA-UTAH] . Bonner Co. : Clark’s Fork, 2100 ft, 17 Aug 1967, HA, LTN, JHL (UCLA 642), 1 M,

1F,6P,4L[UCLA-UTAH]. Bonneville Co.: Swan Valley (5 miS), 5400 ft, 25 Aug 1968, HA, LTN

(UCLA 665), 4 F, 4 L [UCLA-UTAH] . Custer Co.: Challis (12 mi NE), 5200 ft, 13 June 1970, HA

(UCLA 689), 3 IpM (689-15,17,18), 7 lpF (689-11-14,16,19,20), 23 M, 18 F, 18 L [UCLA-UTAH].

Lemhi Co.: North Fork (1 mi S), 3900 ft, 4 Aug 1965, HA (UCLA 655), 1 M, 3 F [UCLA-UTAH].

Madison Co.: Heise Hot Springs, 5200 ft, 26 Aug 1968, HA, LTN (HA-20-68), 1 M, 3 F [UTAH].

Thornton, 4800 ft, 26 Aug 1968, HA, LTN (UCLA 666), 3 M, 18 F, 6 L [UCLA-UTAH].

Montana. Flathead Co.: West Glacier Park, 26 Aug 1959, LTN, 1 F [UTAH]. Granite Co.:

Drummond (2 mi S), 16 Aug 1967, HA, LTN, JHL (UCLA 640), 4 M, 9 F [UCLA-UTAH] . Hall

(6 miS), 4450 ft, 16 Aug 1967, HA, LTN, JHL (UCLA 639), 5 M, 11 F [UCLA-UTAH] . Lake Co.:

Flathead Lake, MSU Biol. Sta., 3000 ft, 12 July 1951, LTN, 2 F [UTAH]. Missoula Co.: Missoula

(27 mi E), 3675 ft, 16 Aug 1967, HA, LTN, JHL (UCLA 641), 9 M, 4 F [UCLA-UTAH] . Ravalli

Co.: Corvallis, 27 June 1948, Jenkins, 1 M, 1 F [UCLA]. Hamilton, 3500 ft, 12 June 1970, HA

(UCLA 688), 9 IpF (688-12-14,16-20), 1 IF (688-15), 16 M, 28 F, 33 L [UCLA-UTAH].

Nevada. Douglas Co.: Glenbrook, 6200 ft, 25 Aug 1915, H.G. Dyar, 1 M, 1 F [USNM] ; same

data, 5 May 1970, HA, RN (UCLA 679), 1 lpM (679-11), 2 IpF (679-12,18), 1 Ip (679-16), 1 L

[UCLA-UTAH].

Oregon. Benton Co.: Corvallis (9 mi N, Berry Creek), 6 June 1959, 5 M [USNM] . Douglas Co. :

Roseburg, 21 Aug 1955, 2 M, 2 F [USNM]. Jackson Co.: Prospect, 26 May 1921, H.G. Dyar, 1 M

[USNM]; same data, 1 June 1921, 1 M [USNM]. Lincoln Co.: Saddleback Mtn., 4 July 1959,

J.C. Dirks-Edmunds, 1 M [USNM]; same data, 11 July 1959, 2 F [USNM]; same data, 25 July

1959, 2 M, 1 F [USNM]; same data, 15 Aug 1959, 1 F [USNM]; same data, 18 June 1960, 1 M,

2 F [USNM]; same data, 16 July 1960, 2 M, 4 F [USNM]; same data, 6 Aug 1960, 1 M, 2

F [USNM]; same data, 27 Aug 1960, 1 M [USNM]; same data, 24 June 1961, 1 M, 2 F [USNM];

same data, 30 June 1961, 1 M [USNM]; same data, 8 July 1961, 3 M, 1 F [USNM]; same data,

9 July 1961, 1 M [USNM]. Marion Co.: Detroit, 21 May 1924, H.G. Dyar, 1 M, 2 F [USNM];

same data, 7 June 1970, HA (UCLA 686), 2 lpM (686-11,16), 6 lpF (686-12-15,17,20), 23 M, 6 F,

30 Contrib. Amer. Ent. Inst., vol. 8, no. 2, 1972

6 L [UCLA-UTAH]. Mill City, 23 May 1924, H.G. Dyar, 1 M [USNM]. Stayton (1 mi N), 200 ft,

7 June 1970, HA (UCLA 687), 2 IpM (687-12,13), 5 IpF (687-11,14,16,17), 1 lp (687-20), 20 M,

60 F, 6 L [UCLA-UTAH]. Tillamook Co.: Rockaway, 27 June 1920, L.P. Rockwood, 25 M

[USNM]. Umatilla Co.: Pendleton (2 mi E), 1070 ft, 18 Aug 1967, HA, LTN, JHL (UCLA

643), 4 M, 14 F, 5 P, 3 L [UCLA-UTAH].

Utah. Weber Co.: Riverdale, 4300 ft, 4 Oct 1970, HA (UCLA 697), 4 lpM (697-11,13,15,17), 1

IpF (697-16), 1 Ip (697-19), 12 M, 36 F, 20 P, 66 L [UCLA]. Trout Springs (3 mi E), 4725 ft,

13 May 1970, LTN (UCLA 685), 6 M, 7 F [UCLA].

Washington. Chelan Co.: Stevens Pass, 25 June 1955, J.N. Belkin, 1 M [UCLA]. Clallam Co. :

Lake Crescent, 3 Sept 1934, A.L. Melander, 2 F [USNM] . Jefferson Co.: Clearwater, 4 June 1934,

CCC Survey, 10 M, 2 F [USNM]. Olympic Natl. Park, Hoh Ranger Station, 4 Aug 1955, LTN, 1 M,

1 F [UCLA]. Quilcene, 20 Aug, R.C. Shannon, 1 F [USNM]. King Co.: Kent, 30 June 1909,

Burke, 2 M, 2 F [USNM]. Seattle, 31 July 1906, Dyar and Caudell, 2 F [USNM]; same data,

17 July 1917, H.G. Dyar, 1 M [USNM]. Lewis Co.: Morton, 10 July 1931, C.M. Gjullin, 1 F

[USNM]. Mason Co.: Belfair, 23 May 1934, CCC Survey, 1 M [USNM]. Hoodsport, 7 July 1920,

H.G. Dyar, 2 M, 4 F [USNM]. Lake Cushman, 27 June 1917, H.G. Dyar, 6 M, 1 F [USNM];

same data, 4 July 1920, 14 M, 4 F [USNM]. Okanagan Co.: Okanagan, 19 July 1920, H.G. Dyar,

1 F [USNM]. Pierce Co.: Ashford,4 Aug 1906, Dyar and Caudell, 3 F [USNM]; same data, 9 June

1921, H.G. Dyar, 3 M, 1 F [USNM]. San Juan Co.: Orcas Island, Mt. Constitution, 7 July 1905,

J.M. Aldrich, 2 M [USNM]. Friday Harbor, 28 May 1906, J.M. Aldrich, 2 M, 1 F [USNM]. Skagit

Co.: Guemes Island, 13 July 1906, J.M. Aldrich, 1 M [USNM]. Skamania Co.: Columbia River,

May 1930, Sam Samson, 8 P, 20 L [USNM] . Snohomish Co.: Stanwood, 5 Aug 1925, A.L. Melander,

1 F [USNM]. Sultan Basin, 2 Sept 1933, CCC Survey, 5 F [USNM]. Walla Walla Co.: Walla Walla,

2-6 July 1922, A.L. Melander, 1 F [USNM]; same data, 18 Aug 1923, 1 F [USNM]. County

Unknown: Canyon Creek, 26 July 1925, A.L. Melander, 8 F [USNM].

NO DATA

1 M, 5 F [USNM].

FIGURES

Distribution of the Varipalpus Group

Mesonotal scale pattern of the Varipalpus Group species

Aedes (O.) laguna; male genitalia and pupa

Aedes (O.) laguna; larva

Aedes (O.) monticola; male genitalia and pupa

Aedes (O.) monticola; larva

Aedes (O.) varipalpus; adult morphology

Aedes (O.) varipalpus; male genitalia and pupa

Aedes (O.) varipalpus; larva

10. Aedes (O.) deserticola; male genitalia and pupa

11. Aedes (O.) deserticola; larva

12. Aedes (O.) sierrensis; male genitalia and pupa

13. Aedes (O.) sierrensis; larva

eT ae oe

|Fig.1

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REFERENCES CITED

Adames, Abdiel J.

1971. Mosquito Studies (Diptera, Culicidae). XXIV. A revision of the crabhole

mosquitoes of the genus Deinocerites. Am. Entomol. Inst., Contrib. 7(2).

154 p.

Anderson, Don L.

1971. The San Andreas Fault. Sci. Am. 225:53-68.

Belkin, John N.

1962. The mosquitoes of the South Pacific (Diptera, Culicidae). Berkeley, Univ.

Calif. Press. 2 vol.

Belkin, John N., S.J. Heinemann and W.A. Page

1970. The Culicidae of Jamaica (Mosquito Studies. XXI). Am. Entomol. Inst.,

Contrib. 6(1). 458 p. (Also published as Inst. Jam., Bull. Sci. Ser. 20)

Belkin, John N. and W.A. McDonald

1956. Aedes sierrensis (Ludlow, 1905), a change in name for the western tree-

hole mosquito of the Pacific Slope (Diptera, Culicidae). Entomol. Soc. Wash.,

Proc. 58:344.

1957. A new species of Aedes (Ochlerotatus) from treeholes in southern Arizona

and a discussion of the varipalpus complex (Diptera, Culicidae). Entomol.

Soc. Am., Ann. 50:179-191.

Belkin, John N., R.X. Schick, P. Galindo and T.H.G. Aitken

1965. Mosquito Studies (Diptera, Culicidae). I. A project for a systematic study

of the mosquitoes of Middle America. Am. Entomol. Inst., Contrib. 1(2):

1-17.

Berlin, O. George W.

1969. Mosquito Studies (Diptera, Culicidae). XII. A revision of the Neotropical

subgenus Howardina of Aedes. Am. Entomol. Inst., Contrib. 4(2). 190 p.

Blaisdell, Frank E. |

1906. Notes and description of the larva of Culex varipalpus Coquillett. Entomol.

News 17:107-109. |

Blanchard, Raphael

1905. Les moustiques. Histoire naturelle et medicale. Paris, de Rudeval. 673 p.

Breland, Osmond P.

1957. Some techniques for collecting tree hole breeding mosquitoes. Mosq. News

17:305-308.

Carpenter, Stanley J. and W.J. LaCasse

1955. Mosquitoes of North America (north of Mexico). Berkeley, Univ. Calif.

Press. 360 p.

Coquillett, Daniel W.

1902. Three new species of Culex. Can. Entomol. 34:292-293.

1906. A classification of the mosquitoes of North and Middle America. U.S. Bur.

Entomol., Tech. Ser. 11. 31 p.

Cupp, Eddie W. and W.R. Horsfall

1969. Biological bases for placement of Aedes sierrensis (Ludlow) in the sub-

genus Finlaya Theobald. Mosq. Syst. Newsl. 1:51-52.

Dyar, Harrison G.

1904a. Notes on the mosquitoes of British Columbia. Entomol. Soc. Wash.,

Proc. 6:37-40.

Arnell & Nielsen: Aedes varipalpus Group 45

1904b. The life history of Culex varipalpus Coquillett. N.Y. Entomol. Soc., J.

12:90-92.

1905a. Remarks on genitalic genera in the Culicidae. Entomol. Soc. Wash., Proc.

7:42-49,

1905b. Illustrations of the abdominal appendages of certain mosquitoes. N.Y.

Entomol. Soc., J. 13:53-56.

1907. Report on the mosquitoes of the coast region of California, with des-

criptions of new species. U.S. Natl. Mus., Proc. 32:121-129.

1917a. The mosquitoes of the mountains of California. Insecutor Inscitiae Mens.

5:11-21.

1917b. The mosquitoes of the Pacific Northwest (Diptera, Culicidae). Insecutor

Inscitiae Mens. 5:97-102.

1918. The male genitalia of Aedes as indicative of natural affinities (Diptera,

Culicidae). Insecutor Inscitiae Mens. 6:71-86.

1919. Westward extension of the Canadian mosquito fauna (Diptera, Culicidae).

Insecutor Inscitiae Mens. 7:11-39.

1920a. The mosquitoes of British Columbia and the Yukon Territory, Canada

(Diptera, Culicidae). Insecutor Inscitiae Mens. 8:1-27.

1920b. The Aedes of the mountains of California and Oregon (Diptera, Culici-

dae). Insecutor Inscitiae Mens. 8:165-173.

1922. The mosquitoes of the United States. U.S. Natl. Mus., Proc. 62:1-119.

1928. The mosquitoes of the Americas. Wash., Carnegie Inst. Wash. (Publ. 387).

616 p.

Dyar, Harrison G. and F. Knab

1906. The larvae of Culicidae classified as independent organisms. N.Y. Entomol.

Soc., J. 14:169-230.

Edwards, Frederick W.

1932. Diptera. Fam. Culicidae. Genera Insectorum 194. 258 p.

Felt, Ephraim P.

1905. Studies in Culicidae. N.Y. State Mus., Bull. 97:442-497, 562-564.

Freeborn, Stanley B.

1926. The mosquitoes of California. Univ. Calif. Publ. Entomol. 3:333-460.

Gardner, Clark F.

1971. Morphology of the mouthparts of larval mosquitoes of the subgenus Och-

lerotatus, genus Aedes. Univ. Utah, Ph.D. dissertation. 216 p.

Hearle, Eric

1926. The mosquitoes of the Lower Fraser Valley, British Columbia, and their

control. Natl. Res. Counc. (Ottawa) Rep. 17. 94 p.

Howard, Leland O., H.G. Dyar and F. Knab

1913. The mosquitoes of North and Central America and the West Indies. Vol. 2.

Plates. Wash., Carnegie Inst. Wash. (Publ. 159). 150 plates. 24 Feb.

1917. The mosquitoes of North and Central America and the West Indies. Vol. 4.

Systematic description (in two parts). Part II. Wash., Carnegie Inst. Wash.

(Publ. 159). p. 525-1064. 31 Mar.

Jenkins, Dale W. and S.J. Carpenter

1946. Ecology of the tree hole breeding mosquitoes of Nearctic North America.

Ecol. Monogr. 16:33-47.

Judson, Charles L., Y. Hokama and J.W. Kliewer

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Entomol. Soc. Am., Ann. 59:1181-1184.

46 Contrib. Amer. Ent. Inst., vol. 8, no. 2, 1972

Knight, Kenneth L. and E.N. Marks

1952. An annotated checklist of the mosquitoes of the subgenus Finlaya, genus

Aedes. U.S. Natl. Mus., Proc. 101:513-574.

Ludlow, Clara S.

1905. A new North American Taeniorhynchus. Can. Entomol. 37:231-232.

1906. Mosquito notes—No. 4. (Continued.) Can. Entomol. 38:132-134.

Lunt, Steele R. and L.T. Nielsen

1971. The use of thoracic setae as a taxonomic tool and as an aid in establishing

phylogenetic relationships in adult female Aedes mosquitoes of North America.

Mosq. Syst. Newsl. 3:102-121.

Matheson, Robert

1929. A handbook of the mosquitoes of North America. Springfield, Ill., Thom-

as. 268 p.

1944. Handbook of the mosquitoes of North America. Ed. 2. Ithaca, Comstock

Publ. Co. 314 p.

McDonald, William A.

1957. The adults and immature stages of Aedes muelleri Dyar (Diptera: Culici-

dae). Entomol. Soc. Am., Ann. 50:505-511.

Myers, Charles M.

1967. Identification and descriptions of Aedes eggs from California and Nevada

(Diptera: Culicidae). Can. Entomol. 99:795-807.

Nielsen, Lewis T., J.H. Arnell and J.H. Linam

1967. A report on the distribution and biology of tree hole mosquitoes of the

western United States. Calif. Mosq. Control Assoc., Proc. Pap., Annu. Conf.

35:72-76.

Nielsen, Lewis T., J.H. Linam, J.H. Arnell and T.J. Zavortink

1968. Distributional and biological notes on the tree hole mosquitoes of the west-

ern United States. Mosq. News 28:361-365.

Nielsen, Lewis T., J.H. Linam and D.M. Rees

1963. New distribution records for mosquitoes in the Rocky Mountain States.

N.J. Mosq. Exterm. Assoc., Proc., Annu. Meet. 50:424-428.

Nielsen, Lewis T. and D.M. Rees

1959. The mosquitoes of Utah—a revised list. Mosq. News 19:45-47.

1961. An identification guide to the mosquitoes of Utah. Univ. Utah Biol. Ser.

12(3). 63 p.

Peyton, E.L.

1956. Biology of the Pacific Coast tree hole mosquito Aedes varipalpus (Coq.).

Mosq. News 16:220-224.

Pratt, Harry D.

1956. A check list of the mosquitoes (Culicinae) of North America (Diptera:

Culicidae). Mosq. News 16:4-10.

Reeves, William C.

1940. Research with Aedes varipalpus (Coq.) the Pacific Coast tree hole mos-

quito. Calif. Mosq. Control Assoc., Proc. 11:39-43.

Richards, Charles S., L.T. Nielsen and D.M. Rees

1956. Mosquito records from the Great Basin and the drainage of the lower Colo-

rado River. Mosq. News 16:10-17.

Rohlf, F. James

1963. Classification of Aedes by numerical taxonomic methods (Diptera: Culici-

dae). Entomol. Soc. Am., Ann. 56:798-804.

Arnell & Nielsen: Aedes varipalpus Group 47

Ross, Edward S.

1943. New and additional Lower California mosquito records (Diptera: Culici-

dae). Pan-Pac. Entomol. 19:86.

Stage, Harry H., C.M. Gjullin and W.W. Yates

1952. Mosquitoes of the Northwestern States. U.S. Dep. Agric., Agric. Handb.

46. 95 p.

Stone, Alan and K.L. Knight |

1956. Type specimens of mosquitoes in the United States National Museum: II,

The genus Aedes (Diptera, Culicidae). Wash. Acad. Sci., J. 46:213-228.

Theobald, Frederick V.

1910. A monograph of the Culicidae or mosquitoes. Vol. 5. Lond., Br. Mus.

(Nat. Hist.). 646 p.

Trebatoski, Alice M. and J.F. Haynes )

1969. Comparison of enzymes of twelve species of mosquitoes. Entomol. Soc.

Am., Ann. 62:327-335.

Zavortink, Thomas J.

1970. Mosquito Studies (Diptera, Culicidae). XVI. A new species of treehole

breeding Aedes (Ochlerotatus) from southern California. Am. Entomol. Inst.,

Contrib. 5(1):1-7.

APPENDIX

Aedes ( ? Ochlerotatus), Jamaican form

As indicated in the section on Systematics there is some suggestion that the

single male (without genitalia), described briefly by Belkin, Heinemann and Page

(1970:143-144) may be realted to the Varipalpus Group. We have therefore pre-

pared a more detailed description here.

MALE. Wing: 3.00 mm. Proboscis: 2.55 mm. Forefemur: 1.90 mm. Medium-

sized species; integument dark brown; white markings on head, thorax, legs; ab-

domen missing. Head: Eyes widely separated above antennae. Decumbent scales

curved, pointed, white on median line and along orbital border, mostly dark lat-

erad of median line, and mixed dark and light posteriorly; orbital line expanded

laterally into patch of broad, white appressed scales, no orbital line below this

patch; postgena with patch of broad, dark scales with white scales below. Forked,

erect scales over dorsal surface, mostly dark. Two pairs of light frontal bristles;

6 pairs of dark orbital bristles. Clypeus dark brown, bare. Proboscis short, 1.38

of forefemur length, entirely dark scaled; a few basal bristles present. Palpus 0.69

of proboscis length, 5-segmented; segment 1 dark scaled; segments 2 and 3 an-

kylosed, making up 0.53 of palpus length, segment 3 expanded apically, both seg-

ments without scales ventrally, remainder dark scaled with narrow white ring at

apex of segment 3, 3 short bristles laterally at apical 0.5 and 0.7 and apex of

segment 3; segment 4 dark scaled, with narrow basal and apical white rings and

48 Contrib. Amer. Ent. Inst., vol. 8, no. 2, 1972

4 or 5 long bristles evenly spaced along lateroventral edge; segment 5 upturned,

dark scaled, with basal white ring and a few white scales apically, a few long

bristles evenly spaced along entire ventral length. Antenna 0.57 of proboscis; torus

enlarged, with white scales on mesal surface; flagellar segments 1-12 with num-

erous long bristles; segment 13 with basal whorl of 6 long bristles; segments 12

and 13 elongate, their combined length about 0.8 of first 11 segments. Thorax:

Mesonotum slightly arched. Mesonotal integument dark brown; partially denuded;

scales all narrow, curved, in indistinct pattern of pale golden on dark brown to

black background, light scales more numerous anteriorly, in posterior fossal area,

outer posterior dorsocentral area and around prescutellar space. Acrostichal, dor-

socentral, prescutal and supraalar bristles present, well developed. Paratergite with

white scales, narrow, curved anteriorly, broad posteriorly; with mixed light and

dark bristles. Scutellum dark brown, lobes with patches of narrow, curved white

scales and about 8 long, dark bristles and several smaller bristles. Postnotum dark

brown, bare. Pleural integument brown, lighter around spiracle and at wing base.

Apn not enlarged. Pleural bristles on apn, ppn, ppl, psp, pra, stp, upper mep,

lower mep. Apn with patch of mostly erect, broad white scales; ppn covered pos-

teriorly with broad white appressed scales and a few dark scales; patches of broad,

white appressed or semierect scales on ppl, psp, ssp, pra below knob; 2 patches

on stp, 1 on upper margin and 1 posteriorly at level of midcoxa; patch on upper

mep extending ventrally to about middle of sclerite; metameron, hypostigial area

and meron without scales. Legs: Rather short, forefemur about 1.25 height of tho-

rax (mesonotum to apex of midcoxa). Hindcoxa subequal to midcoxa, its base

well below upper margin of meron. Forecoxa with white scales basally and api-

cally, dark scales between; midcoxa and hindcoxa with white scales. Trochanters

with white scales. Femora mostly dark scaled, with narrow basal and apical white

rings and scattered white scales, white scales forming streak on basal ventral and

dorsal surfaces of forefemur to near apex and basal ventral surface of midfemur

and hindfemur to near apex. Foretibia dark scaled, with a few apical white scales;

midtibia dark scaled, with a few basal and apical white scales. Hindtibia missing.

Foretarsus dark scaled, with narrow basal white ring on base of segment 1, nar-

row basal white ring and white apical dorsal patch on segment 2; midtarsus dark

scaled, with narrow basal ring and apical white patch on segments | and 2 and

narrow basal white ring on segment 3. Claws of foreleg and midleg enlarged, un-

equal, with acute subbasal tooth. Wing: Entirely dark scaled. Haltere: Light brown,

with white scales on upper part of knob. Abdomen: Tergite I with few white

scales mesally; laterotergite with small, dense patch of white scales. Remainder

of abdomen missing.

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Contributions

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American Entomological Institute

Volume 8, Number 3, 1964

MOSQUITO STUDIES (Diptera, Culicidae)

XXVIII. The New World species formerly placed in

Aedes (Finlaya). By Thomas J. Zavortink

MOSQUITO STUDIES (Diptera, Culicidae)

XXVIII. THE NEW WORLD SPECIES FORMERLY

PLACED IN AEDES (FINLAYA ,

by

Thomas J. Zavortink’

CONTENTS

INTRODUCTION 3

SYSTEMATICS . . , 3

MEDICAL IMPORTANCE ae 4

KEYS TO GENERA AND SUBGENERA 4

TAXONOMIC TREATMENT . . ENS Oa eet ree ee re ee fey rents

Genus Aedes, Subgenus Pro ey eT a Tee ay ane ene ere

Keys to Groups Debra is EN tee caiga ies Line seeemabinaetene Tinea itt ea! Tat

Triseriatus Group. i?

Keys to Species ... 19

1. Aedes (P.) hendersoni 20

D> ete ere a ahh a he eal ge Ce gos vans co ae se a eee

S. eR) BrSeHG TS. 2) eR gis ie I ees Nan de ee ie Ghee

Zoosophus Group. . Ag naY ae.) “ire RRC tah aoe ener a

4, Aedes (P.) zoosophus IY Se segle, Sia Tp FSH ek eee Rt EA age de ke

Komi ahigaee 236 oie sk ae 5 taotine’ hea ee ating orbs Oak CAE wt arian eee

Keys.to,Species.. .... eames Staak te, Ss eal em Be aT one's ahaa a lee eg an iain ee

5. Aedes (P.) puréeria, Syernae . escaasay lh Weed, | nee ck Dice ike amare abe: Pao cage

6. AvdestP PROM fo yo te ied By a tahicks (407s Re ates Shee

Fn, Aedesd PJ BCR oo iat, eA SOR ee ee

8. Aedesd Pd chionetiiis ides nek Gvarmcomeitie wine Serta

9. Aedes (PF Paieoseuiittvinc: i we noc Unies tae oes Cees

LO. Aedest? Jeondiae cosets aah Sree hee Te Daten aie oS

KnabiGroup . . 1927 tad pikean Cees gad Paraben te ee, Sa

11. Aedes (P.) bhated vinges” tecabiay hwapanmis rab LARC ae ennai ies, “em

Subgenus Kompia . . sd cura gu ea ay edae tenet ache rs are ar Where ean

12. Aedes (K.) purpureines yr etsy pene Sie aie Plan wie A ee ee

Subgenus Abraedes . . . og 4d peeve ved satan eS “legen ate Se oe A gi, ae

13. Aedes (Abr.) papago . ge Sy Sa mare eee PR erie ee ae

‘Contribution 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.

* Department of Zoology, University of California, Los Angeles, California 90024.

Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

_ Subgenus Aztecaedes te

14. Aedes (Azt.) ramirezi

Subgenus Gymnometopa . :

15. Aedes (G.) diovan ;

Subgenus Ochlerotatus . ;

Pulchritarsis Section, Muelleri Eronp :

16. Aedes (O.) muelleri .

Atropalpus Section .

Keys to Groups

Atropalpus Group

Keys to Species :

17. Aedes (O.) enariius

18. Aedes (O.) Bea

Fluviatilis Group . .

19. Aedes (O. ye

Genus Haemagogus, Subgenus Conoposteaus :

Keys to Species :

20. Haemagogus (C. ) ele ee

21. Haemagogus (C.) sp., Large © ombicn cat

22. Haemagogus (C.) sp., Peruvian highland form

23. Haemagogus (C.) leucophoebus

24. Haemagogus (C.) leucocelaenus

25. Haemagogus(C.) clarki. . .

26. Haemagogus (C.) sp., Small Colombian for

27. Haemagogus (C.) sp., Peruvian lowland form

REFERENCES CITED

FIGURES

SYSTEMATIC INDEX

Zavortink: New World Aedes 3

INTRODUCTION

The present revision includes the majority of the species of New World Aedes

(Finlaya) exclusive of the Terrens Group, as well as Aedes (Abraedes), Aedes

(Kompia) and Aedes (Ochlerotatus) muelleri. A total of 23 named species, known

in all stages, and 4 unnamed forms, known by 1 or more females, is treated.

In all, 22,235 specimens, 4,164 males, 4,967 females, 7,542 larvae, 5,562 pupae

and 2,424 individual rearings (1,497 larval, 767 pupal, 160 incomplete) have been

examined.

The methods of study and presentation, as well as the terminology and ab-

breviations, follow, in general, Belkin (1962); a few additional terms for the meso-

scutal markings and details of the genitalia follow Berlin (1969) and Schick (1970a),

respectively.

I am indebted to John N. Belkin for reading the manuscript and suggesting

certain improvements; Harold C. Chapman, George B. Craig, Jr., Pedro Galindo,

Kenneth L. Knight, Lewis T. Nielsen, George F. O’Meara, L.L. Pechuman, John

F. Reinert, Paul L. Rice, Alan Stone, Floyd G. Werner, and Michael J. Zavortink

for loans of museum specimens or gifts of live mosquitoes; Willis W. Wirth for

information on the type of Finlaya nigra; Abdiel J. Adames, J. Hal Arnell, Lewis

T. Nielsen and Robert X. Schick for stimulating discussions; Sandra J. Heinemann

and William A. Powder for help in the preparation and rearing of specimens;

Sharon Burmeister, L. Margaret Kowalczyk and Nancy L. Martsch for prepara-

tion of many of the preliminary drawings and L. Margaret Kowalczyk for prep-

aration of all of the final illustrations; Claire M. Price for typing a portion of ©

the preliminary draft; and, finally, Caryle Stallard for typing the remainder of

the preliminary draft, providing editorial assistance and preparing the text copy

for lithoprinting.

SYSTEMATICS

I am removing all New World Aedes that have been placed in the subgenus

Finlaya from that subgenus and assigning them to other subgenera or genera, as

follows: the Triseriatus, Zoosophus, Kompi and Knabi Groups treated in the pres-

ent paper and the Terrens Group treated by Schick (1970a,b) are placed into

the subgenus Protomacleaya of Aedes; ramirezi and mediovittatus are placed into

the monotypic subgenera Aztecaedes and Gymnometopa of Aedes, respectively;

the Atropalpus and Fluviatilis Groups are placed into the subgenus Ochlerotatus

of Aedes; scutellalbum and upatensis are transferred to Ochlerotatus, where they

are related to milleri Dyar, 1922 and lepidus Cerqueira and Paraense, 1945, re-

spectively, and are not considered further because of lack of material; and, the

Leucocelaenus Group is placed into the subgenus Conopostegus and transferred

to Haemagogus. The present paper also treats the monotypic subgenera Kompia

and Abraedes of Aedes and Aedes (Ochlerotatus) muelleri.

In hope of making the keys that follow more useful, I have included in them,

where possible, the following additional groups of New World container-breeding

Aedes: the subgenus Howardina, the Terrens Group of Protomacleaya, the Vari-

palpus Group of Ochlerotatus, and Aedes (Stegomyia) aegypti. Additional infor-

mation on these taxa may be found in the revision of Howardina by Berlin (1969),

4 Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

the revision of the Terrens Group by Schick (1970a,b), the revision of the Vari-

palpus Group by Arnell and Nielsen (1972), and, for aegypti, ‘““The Culicidae of

Jamaica” by Belkin, Heinemann and Page (1970).

MEDICAL IMPORTANCE

Aedes triseriatus is known to be a vector of the LaCrosse strain of California

encephalitis in Ohio and Wisconsin, Aedes fluviatilis may have been a vector of

sylvan yellow fever in Brazil on 1 occasion, and species of Haemagogus (Con-

opostegus) have been found naturally infected with yellow fever in Brazil and

Colombia and with the arbovirus Una in Brazil. In addition, laboratory experi-

ments have shown Aedes atropalpus, Aedes fluviatilis, Aedes triseriatus and Haema-

gogus leucocelaenus capable of transmitting 1 or more of the following: eastern

equine encephalomyelitis, western equine encephalomyelitis, Venezuelan equine

encephalomyelitis and yellow fever. Additional details on natural and artificial

infections can be found in the bionomics section of the 4 species just named

and the large and small Colombian forms of Haemagogus (Conopostegus).

In addition, it should be stressed that most of the species included in the pres-

ent revision are known to be anthropophilic and many are capable of breeding

in natural or artificial containers in domestic or quasi-domestic situations, and,

as a consequence, all these are of potential importance as vectors of arboviruses.

KEYS TO GENERA AND SUBGENERA

ADULTS

- Mesoscutum with moderately broad to broad flat scales; mesoscutum with

metallic coppery, green, blue or violet scales or pleuron with 2 very

conspicuous, nearly vertical bands of scales. . . . . . . Haemagogus

Mesoscutum with narrow curved scales; mesoscutum never with metallic

coppery, green, blue or violet scales; pleuron usually without vertical

bands of scales, rarely with a single conspicuous vertical band. . . . .2

2(1). Postspiracular area without bristles (12. purpureipes) . . . Aedes (Kompia)

Postspiracular area with bristles . RL eee ©

3(2). Abdominal tergites VI and VII with a pair of large, bristly, scaleless areas;

sternopleuron with 2 narrow, diagonal, median lines of silver scales (13.

papago) .. ‘ . . . Aedes (Abraedes)

Abdominal tergites VI and VIL completely scaled: sternopleuron with a

single narrow, median, diagonal or horizontal line of scales, a single med-

ian patch of scales, or nearly entirely covered with scales . .... .4

4(3). Mesoscutum with a very narrow white-scaled median acrostichal line that

is continued as a median prescutellar line through prescutellar space to

scutellum; EY bare; tarsi entirely dark scaled (14. ramirezi) .

5 Aedes (Aztecaedes)

Wecoecutun. never arn tino ‘ight- edled median acrostichal and med-

5(4).

6(5).

7(6).

8(7).

9(8).

10(9).

Zavortink: New World Aedes 5

ian prescutellar lines; paratergite with scales and/or tarsi with 1 or more

Lahti AAS sine ars) oebiwisinh iluntysts «ase te epee SB epeemaue id wae 8D

Clypeus with scales; scutellum with broad flat silver-white scales on all lobes

forming a complete transverse band (aegypti; see Systematics) :

: . Aedes ( Steconipia)

Clypeus without scales: scutellum “without eae flat white scales on all

lobes: fonming. ancomipete transverse. band, 2° fan, aie lek Ts UD

Sternopleuron without scales at base of prealar knob; hindtibia with con-

spicuous patch of white scales about 0.3-0.4 distance from base (15.

mediovittatus) . . . . Aedes (Gymnometopa)

Sternopleuron with scales at base of prealar knob: hindtibia without patch

of white scales about 0.3-0.4 distance from base. . . .......7

Laterotergite of abdominal segment I without scales or outer surface of

hindcoxa bare (Atropalpus Section) . . . in part Aedes (Ochlerotatus)

Laterotergite and outer surface of hindcoxa with scales . . ..... .8

Hindtarsal segments 2 and 3 with conspicuous broad white apical band

(Varipalpus Group—in part Pulchritarsis Section; see Systematics) .

. in part Aedes ( Ochlerotatus)

Tian, seoments. 2D asia 3 stan dav scaled or 1 or both with light

Wasa) BAGG oe tadtis reves Fats ear ee es Deas Fes ee

Hypostigial area with patch of white scales and hindtarsus entirely dark

scaled in muelleri—in part Pulchritarsis Section) .

. in part Aedes ( Ochlerotatus)

Hypostigial area s usually cee if scaled then hindtarsus with at least 1 light

Dall. Laer « fo aft «had CagtReOte anaes ae 4° 8

Vertex of head usually with narrow curved scales and numerous erect scales;

mesepimeron usually with a single large patch of scales covering most

of sclerite; tarsi varied, but light scales, if present, not restricted to band

at base of segment 1 of alllegs. . . . . . . . Aedes (Protomacleaya)

Vertex of head with broad flat scales except along midline and with few

or no erect scales; mesepimeron usually with 1 or 2 small patches of

scales; if with a single large patch then only tarsal segment | of all legs

with a light basal band (see Systematics). . . . . .Aedes (Howardina)

FEMALE GENITALIA

Cercus obliquely truncate in lateral aspect and with 2-4 strongly developed

setae at apex; tergite IX well developed, moderately to strongly sclero-

tized, rather narrow anteriorly and with a pair of narrow elongate pos-

terior extensions so that it is more or less U- or V-shaped . Haemagogus

Cercus not obliquely truncate in lateral aspect and without strongly de-

veloped setae; tergite IX weakly to moderately developed and sclero-

tized, ‘usually not distinctly U-or Veshaped .... 6 ws ag eee

2(1).

3(2).

4(3).

5(4).

6(5).

7(5).

8(7).

9(8).

10(8).

Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

Insula without setae; apical margin of sternite VIII deeply emarginate be-

tween a pair of more strongly sclerotized submedian lobes (aegypti; see

Systematics)... . ... . . Aedes (Stegomyia)

Insula with at least 1 bait of peevicnous elongate ae apical margin

of sternite VIII straight or shallowly emarginate, without differentially

sclerotized subiiedian lobes 9 se ee ee te ee ee

Tergite VIII with scales absent or few (Varipalpus Group—in part Pulchri-

tarsis Section; see Systematics). . . . . . in part Aedes (Ochlerotatus)

Tersite Vill wath! numerous’scalés 545 ee eer eee eh

Tergite IX reduced to a pair of separate, weakly sclerotized, dorsolateral

plates without setae; postgenital plate with a pair of setae that are as

strongly developed as setae of cercus (Atropalpus Section) . .

; . in part Aedes ( Ochlerotatus)

Tergite 1x not reduced to a “pair ‘of dorsolateral plates and usually with

at least 1 pair of setae; postgenital plate with setae that are not as strong-

ly developed as*setacor cereus: 2 Vr Fr Eee er eee LS

Aifial places preser B eh RO ei. og He LOS ee PG

Atrial plates absent. #)

Postgenital plate moderately long and broad; cercus with numerous scales;

basal portion of sree duct not sclerotized (15. mediovittatus)

seats . Aedes (Gymnometopa)

Postgenital plate oie and narrow; cercus without scales; basal portion of

spermathecal duct sclerotized . . . . . . . . Aedes (Protomacleaya)

Tergite IX lobe with 11-17 moderately developed setae; insula with same

degree of sclerotization and Haleass as sigma (12. purpureipes)

. Aedes (Kompia)

Teroite 1X be with 9 or oe setae or inet much more strongly scle-

rOtlged and Digmientcd Milan sigma 6 7%. Te ee. a

Sternite VIII with relatively few conspicuous scales . ....... .9

Sternite VIII with very numerous conspicuous scales ...... . 10

Insula weakly sclerotized and pie iied tergite IX lobe with 1 seta; cercus

with scales (13. papago) . . . fk eges (Abraedes)

Insula strongly sclerotized and pigmented: tergite IX lobe with 5-9 setae;

cercus without scales (14. ramirezi). . . . . . . .Aedes (Aztecaedes)

Postgenital plate with moderately to strongly developed basal median apo-

deme; tergite VIII with conspicuous setae on disc and without a dense

band of outstanding scales distally (16. muelleri—in part Pulchritarsis

DOCHOM). uss . . . in part Aedes (Ochlerotatus)

Postgenital plate without a Basal ‘median apodeme; tergite VIII usually with-

out conspicuous setae on disc and usually with a dense band of out-

standing scales distally (see Systematics). . . . . .Aedes (Howardina)

2(1).

3(2).

4(3).

5(4).

6(5).

7(6).

8(7).

9(8).

Zavortink: New World Aedes 7

MALE GENITALIA

Sternomesal surface of sidepiece with long broad striated scales; paraproct

with striated knob at apex .. . sii Haewiagogus

Sternomesal surface of sidepiece without jong ‘Broad striated scales; para-

proct with. |, or. rately more apical tect ae eis et yee ee

Aedeagus with apical and lateral teeth; cercal setae absent; paraproct with

long sternal arm (aegypti; see Systematics) . . . . . Aedes (Stegomyia)

Aedeagus without apical or lateral teeth; cercal setae present; paraproct

without. sternal: Gran. igo peso ie aa ocean ete geet

Claspette represented by a small basal lobe or large plaque on sternomesal

margin of sidepiece and never bearing a single long, strong, curved fila-

ment (see Systematics). . . . . . . .Aedes (Howardina)

Claspette developed as a columnar stem projectine caudad mesad of side-

piece and bearing a single, long, strong, curved filament ..... .4

Proctiger unusually long in dorsal aspect, the basolateral sclerotization nearly

horizontal (13. papago) . . . . Aedes (Abraedes)

Proctiger not unusually long in dorsal Aspect, the ‘basolateral sclerotization

HACGMY VerUlCal 456 SMe ee LS AMIR A RIES pee em, | MS BR LN ae

Tergite IX without sclerotized lobes and without setae (Atropalpus Section)

. in part Aedes (Ochlerotatus)

Tergite IX ‘with small to ae sclerotized lobes bearing 2 or more setae . .6

Claspette stem with 2 strongly developed, elongate setae arising on dorsal

surface distally (14. ramirezi) . . . . . . .Aedes (Aztecaedes)

Claspette stem without strongly developed setae PaAaie CM Rade cs hs cag OE OM |

Claspette filament with a distinct, large, retrorse angle on convex side;

spiniform of clasper long, 0.5-0.7 length of clasper, and basal tergomesal

area of sidepiece with a single, long, strong, differentiated seta (15. medio-

vittatus) . . . . . . Aedes (Gymnometopa)

Claspette fitenient autliont distinct Foun angle; spiniform shorter than

0.4 of clasper length and/or basal tergomesal area of sidepiece without

a lone: strong; differentiated’ seta’ 2) ho. <a ee a eg tee re Rl Pd

Aedeagus distinctly expanded distally in dorsal aspect; tergite IX lobe with

8-17 setae in several series (12. purpureipes) . . . . . Aedes (Kompia)

Aedeagus not expanded distally in dorsal aspect; fergite IX lobe with 2-

10’ setae ina single or partially double series (20. eee ae Pee i 9

Basal tergomesal area of sidepiece with clump of few to very numerous

weakly to moderately developed setae; clasper without setae on outer

surface near apex; spiniform of clasper long, 0.3-0.6 length of clasper

4 . Aedes (Protomacleaya)

Basal tereomiedal area of Ridepicce ae 1 or more long, strong, differenti-

ated setae; clasper with 1 or 2 setae on outer surface near apex; spini-

form of claspershort, 072-0.3 tenet of clasper 2". ere, Se

10(9).

2(1).

3(2).

4(3).

5(4).

6(5).

7(6).

8(7).

Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

Tergal surface of sidepiece with numerous scales among the shorter setae;

aedeagus usually broadest in middle and more or less barrel-shaped in

dorsal aspect (16. muelleri—in part Pulchritarsis Section)

. in part Aedes ( Ochlerotatus)

Tergal mupiace ‘of sidepiece without scales or with very few scales among

the shorter setae; aedeagus usually broadest at base and slightly tapered

distally in dorsal aspect (Varipalpus Group—in part Pulchritarsis Section;

see Systematics) . . ...... . . . in part Aedes (Ochlerotatus)

PUPAE

[Aedes (Howardina), Varipalpus Group of Aedes (Ochlerotatus)

and Terrens Group of Aedes (Protomacleaya) not included ]

Float hair (1-I) usually displaced mesad and 1-II usually displaced laterad,

so that distance between alveoli of float hairs is usually less than 0.6

distance between alveoli of hairs 1-II .... . . . Haemagogus

Distance between alveoli of float hairs usually greater than 0.7 distance

Dei Weer alveoli iat belie invent os ant wiles taaitoe - ted

Hair 6-C longer and stronger than 7-C aeaiae see Systematics) ...

<es . Aedes ( (Stegomyia)

Hair 6-C shorter and weaker than ue eal or

Paddle with long filamentous marginal spicules (13. papago) . .

F Aedes s (Abraedes)

Paddle without lone filamentous marginal spicules .

Tergites II-VI or VII with a long, strong, irregular transverse ridge slightly

caudad of level of hair 0; midrib of paddle conspicuous only basally

(14. ramirezi) ... . . .Aedes (Aztecaedes)

Tergites II-VI or VII without: a dcansyetse: ridge caudad of level of hair 0

or midrib of paddle conspicuous to or nearly to apex of paddle . . . .5

Hair 6-VII strongly developed, longer and stronger than 6-VI and 9-VII

(15. mediovittatus) . . anid Aedes (Gymnometopa)

Hair 6-VII weakly to moderately. developed, weaker and shorter than 6-

Vivand/or-9-Vib eo atte ber ; omer oe 6G

Paddle as broad as or broader than long (Atropalpus Section) :

. in part Aedes ( Ochlerotatus)

Paddle disinedy bangs (iad bread Biet, betes POR erotas. . 47

Hair 5-IV,V short, 0.4-0.7 length of corresponding tergite (12. purpureipes)

iy . Aedes (Kompia)

Hit Ss IV Vv moderately loti " one 1. 0- Ja 2 feneth of Cage aie tergite

Hair 1-V or VI to VII much more weakly developed than hair 1 of immed-

iately preceding segments; 6-IIJ-VI relatively strongly developed (16. muel-

peas

a

4(1).

5(4).

6(5).

7(4).

8(7).

Zavortink: New World Aedes 9

leri—in part Pulchritarsis Section). . . . . in part Aedes (Ochlerotatus)

Hair 1-V or VI to VII not conspicuously weaker than hair 1 of immedi-

ately preceding segments; 6-III-VI weakly to moderately developed

in part Aedes (Protomacleaya)

LARVAE

Hair 3-VII strongly developed, very long, usually single (single or double) . 2

Hair 3-VII weakly to moderately developed, short, single to multiple. . .4

Pecten of siphon strongly arcuate dorsad distally; comb scales with | or

2 long and usually 1-3 shorter unfringed spines from a large basal plate

(15. mediovittatus) . ... . . . . . Aedes (Gymnometopa)

Pecten of siphon more or less ahr tre comb aang with a single fringed

or serrate apical projection from a small base ..........3

Hair 9-III-V as long and strong as or longer and stronger than hair 7 of

corresponding segment; 12-I absent (see Systematics) .

. Aedes ( Newivdina)

iit 9- IIL-V aes? mal wens ee fair 7 of corresponding segment and/

or TZ present 2 0. Pl ae ne PP ees pe ee eres

Ventral brush (hair 4-X) arising from a weakly to strongly sclerotized boss

Ventral brush artine *ivomy ame A ee ed ee ne

Boss of ventral brush weakly sclerotized; siphon without acus; labial plate

subquadrate (13. papago). . . . . . . . . Aedes (Abraedes)

Boss of ventral brush strongly alneanineds eerie with distinct acus; labial

plate distinctly netrowed apicallyiiuesiity) scape 2 othe, bere. ee}

Integument of thorax and abdomen with conspicuous simple spicules; pecten

of siphon relatively long, more or less straight; comb scales with long

unfringed apical spine (12. purpureipes) . . . . . . . Aedes (Kompia)

Integument of thorax and abdomen without conspicuous spicules; pecten

of siphon short, conspicuously arcuate dorsad distally or oblique; comb

scales fringed (14. ramirezi). . . . . .. . . . Aedes (Aztecaedes)

Comb scales with long median spine and strong basal spinules; ventral brush

with 5 pairs of hairs (aegypti; see Systematics). . . . Aedes (Stegomyia)

Comb scales usually fringed; if with long median spine, then without strong

basal spinules and ventral brush with more than 5 pairsof hairs . . . .8

Comb scales with long unfringed apical spine; most caudal hair of ventral

brush (4a-X) short; ventral brush with 7 pairs of hairs (16. muelleri—in

part Pulchritarsis Section) Oi? . . in part Aedes (Ochlerotatus)

Comb scales fringed, the apical element of the fringe rarely produced into

moderately long spine; most caudal hair of ventral brush usually long,

10 Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

if short then ventral brush with only 6 pairsof hairs . ...... .9

9(8). Ventral brush with 6 pairs of hairs and the more caudal hairs (4a-c-X)

single or double ee Group—in part Pulchritarsis Section; see

Systematics). . . . . . . in part Aedes ( Ochlerotatus)

Ventral brush varied, but never with 6 pairs of hairs and the more caudal

hairs single or double eee aon mirth Pele eis SacetekO

10(9). Hair 5-I]-IV moderately to strongly developed; comb scales not irregularly

cleft and distal pecten teeth not strongly detached :

eae Aedes (Porpeneeava )

Hat efL- IV weakly ‘developed: comb scales irregularly cleft or distal pec-

ten teeth ai detached si as cal Section) . .

wae : . in part hiedes ( Bieiierovitus)

TAXONOMIC TREATMENT

Genus AEDES Meigen

1818. Aedes Meigen, 1818:13. TYPE SPECIES: Aedes cinereus Meigen, 1818, Europe; mono-

basic.

For complete synonymy see Stone, Knight and Starcke (1959).

Subgenus PROTOMACLEAYA Theobald

1904. Gualteria Lutz, 1904a:13, 1904b:4. TYPE SPECIES: Gualteria oswaldi Lutz, 1904,

Brazil; first of 2 included species, selection of Brunetti (1914:61). Preoccupied by

Gualteria Agassiz, 1872.

1907. Protomacleaya Theobald, 1907:253-254. TYPE SPECIES: Culex triseriatus Say, 1823,

United States of America; monobasic and original designation.

Protomacleaya of Theobald (1907:253-254; 1910:250, in part).

Aedes (Gualteria) in part of Lutz (1904a:13); Dyar (1918:79); Vargas (1950:62-63); Vargas

and Downs (1950:161-172).

Aedes (Finlaya) in part of most authors.

FEMALES. Dark scales of palpus, proboscis, legs, wing and abdomen with blue

or violet reflections. Head: Eyes narrowly separated above antennae, the result-

ing space with light narrow curved scales. Integument yellow to tan or brown

to black. Frontal bristles strongly developed, 1-3 pairs. Orbital bristles relatively

few, mesal 2-5 pairs strongly developed, lateral 4 or 5 pairs moderately devel-

oped. Vertex and occiput with numerous narrow curved decumbent scales and

numerous erect scales. Orbital line of light narrow curved scales present. Dorso-

lateral, lateral and ventral surfaces with broad flat scales. Clypeus moderately large,

bare. Proboscis relatively stout, longer than to subequal in length to forefemur;

entirely dark scaled; with a few basal bristles. Palpus moderately long, about 0.22-

0.29 of proboscis; 3- or 4-segmented, segment 4 very small to minute when pres-

ent; segments 1-3 with bristles; entirely dark scaled. Antenna subequal in length

Zavortink: New World Aedes 11

to proboscis or slightly longer; torus frequently without scales, scales small and

inconspicuous when present; flagellar segment 1 longer than 2, usually slightly

swollen, with small scales; flagellar segments 2-13 usually with 6 moderate bristles

in basal whorl; each of flagellar segments 3-13 slightly longer than preceding.

Thorax: Integument yellow to tan or brown to black. Acrostichal and dorso-

central bristles in complete well developed rows or reduced in number and extent

and then sometimes totally absent from disc; prescutellar and supraalar bristles

numerous and well developed; humeral, lateral prescutal and posterior fossal bris-

tles varying from numerous and well developed to absent; | parascutellar bristle

present. Scutellum with 4-10 strong bristles on midlobe, 3-10 on lateral lobe.

Mesoscutum completely covered with narrow curved scales except for anterior

inner dorsocentral and median and lateral prescutellar bare spaces; pattern varied.

Scutellum with narrow curved and/or broad flat scales on midlobe, narrow curved

scales on lateral lobe; color of scales varied. Paratergite broad, with patch of light

scales. Apn not enlarged. Ppn separated from mesoscutum by strong suture. Meron

large. Pleuron with bristles on apn, ppn, ppl, psp, stp, pra and upper mep; stp

bristles in long row from psp bristles to below lower margin of mep; ssp and low-

er mep bristles absent. Apn with broad and/or narrow curved scales, neither type

closely appressed; ppn nearly entirely covered with scales, these narrow curved

and/or broad flat; broad scales in patches on ppl, stp below pra, stp caudad of

ssp area, stp cephalodorsad of midcoxa, and in a single large patch extending

from upper bristles nearly to lower margin on mep; additional patch usually on

ssp and sometimes on pst, psp and metameron; pcx and hypostigial scales ab-

sent. Legs: Hindcoxa subequal in size to or smaller than midcoxa, its base far

below upper margin of meron. Legs moderately long, the forefemur about 1.18-

1.40 times distance from top of thorax to tip of midcoxa. Forecoxa with light

or light and dark scales on outer surface; midcoxa and hindcoxa with light scales

on outer surface. Forefemur and midfemur dark scaled with light scales in at

least basal 0.5 of posterior and sometimes dorsal and/or ventral surfaces and some-

times at base on anterior surface; hindfemur light scaled in at least basal 0.5,

dark scaled in apical portion; light scales on posterior and/or ventral surfaces of

all femora sometimes extending to apex. Knee spots small to large, white to sil-

ver; largest on hindleg, sometimes absent on foreleg. Tibiae basically dark scaled,

sometimes with basal light patch or ring or light streak on posterior surface. Tarsi

varied. Claws of foreleg and midleg with acute submedian tooth; claws of hind-

leg varied. Wing: Entirely dark scaled or with light scales at base of costa and/or

vein R. Plume scales present on dorsum of veins Rs, R,+3, R2, R3, M basad of

furcation, and less conspicuously on apical portion of 1A. Haltere: Knob light

and/or dark scaled. Abdomen: Tergite I with numerous scales middorsally. Lat-

erotergite with large patch of white or silver scales. Segments VI and VII com-

pletely scaled. Without outstanding scales. Ornamentation varied.

FEMALE GENITALIA (figs. 5,14). Segment VII: Tergite semielliptical in shape,

length along midline about 0.55-0.65 length of tergite VII; distal 0.65-0.75 with

scales and bristles. Sternite long, about 1.2-1.4 length of tergite, much broader

distally than basally; distal margin broadly emarginate; all but narrow basal and

lateral portions with scales or bristles, the latter predominating; bristles more

numerous along midline and distal margin, largely weakly developed and short,

but some moderately developed and elongate. Tergite 1X: Deeply notched apically,

maximum length about 0.45-0.55 of tergite VIII; moderately sclerotized; apical

lobe with 1-7 fine short bristles. Jnsula: Weakly sclerotized; connected to sigma;

12 Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

with 2-5 pairs of moderately developed bristles. Cercus: Long, length of dorsal

edge 0.85-0.95 length of tergite VIII; compressed; apex rounded in lateral view;

bristles numerous apically, the largest moderately developed; scales absent. Post-

genital Plate: Long and narrow, length 0.65-0.80 of tergite VIII; index about 2.8-

3.3; apex rounded in ventral view; distal portion with numerous weakly developed

bristles; basal median longitudinal apodeme strongly sclerotized. Cowl: Strongly

sclerotized. Atrial plates moderately large, strongly sclerotized. Sigma: Continuous

with cowl; weakly to moderately sclerotized. Basal portion of spermathecal duct

strongly sclerotized. Spermathecae 3, strongly sclerotized, more or less spherical,

1 slightly larger than others.

MALES. Essentially as in females except for sexual characters. Head: Proboscis

relatively stout. Palpus subequal in length to or slightly longer than proboscis;

5-segmented; segments 2 and 3 ankylosed and long, making up 0.62-0.67 length

of palpus; segment 4 short, 0.16-0.18 length of palpus; segment 5 short, 0.12-

0.15 length of palpus; palpus slender except for swollen apex of segment 3

and swollen segment 4; apex of segment 3 upturned, segments 4 and 5 droop-

ing; apex of segment 3 and all of segments 4 and 5 with long bristles; usually

completely dark scaled. Antenna subequal in length to or slightly shorter than

proboscis; torus much enlarged, bare; flagellum strongly plumose, segments 1-12

with very numerous long bristles; flagellar segment | slightly elongate, with scales;

flagellar segments 12 and 13 elongate, subequal in length or penultimate longer,

the 2 combined nearly equal to total length of first 11 segments. Legs: Claws of

foreleg and midleg enlarged, unequal; larger claw with blunt submedian tooth and

with or without an acute basal external tooth; smaller claw with acute submed-

ian tooth. Claws of hindleg small, varied. Abdomen: Apical segments and geni-

talia not conspicuously bent ventrad.

MALE GENITALIA. Predominantly various shades of straw-yellow to brown

with apex of paraproct dark brown to black. Segment VIII: Tergite long; 0.78-

0.98 length of sternite; exserted, apical 0.50-0.70 scaled. Segment IX: Well de-

veloped; middorsal portion of tergite weakly to moderately sclerotized, at or cau-

dad of level of base of sidepiece; tergite lobes prominent, narrowly to broadly

connected middorsally, each with 1-9 moderately to strongly developed setae in

1 series; sternite large, with 2-8 setae distally. Sidepiece: Well developed, usually

subcylindrical to subconical; mesal surface membranous from base to apex; basal

tergomesal area slightly to conspicuously swollen, with few to numerous weak-

ly to moderately developed setae; apical lobe absent; median sternomesal scle-

rite absent to well developed and then with a conspicuous tuft of setae; sterno-

mesal surface without long broad striated scales; dorsal, lateral and ventral sur-

faces with numerous scales and bristles; bristles of ventral surface not specialized.

Claspette: Well developed; stem short, straight in dorsal aspect; spiculose; with 2-

6 weakly developed setae; filament long, subterete to distinctly flattened apically,

curved or angled dorsad, with or without ridge on convex side, without retrorse

barb. Clasper: Simple, short, swollen subbasally; median portion straight, with

longitudinal ridges or wrinkles; basal portion spiculose; apical portion with 1 or

2 fine setae on inner surface, none on outer surface; apical spiniform long, 0.35-

0.50 length of clasper. Phallosome: Aedeagus moderately large, without teeth;

base slightly bulbous, remainder subparallel-sided with truncate apex in dorsal as-

pect. Proctiger: Strongly developed; not unusually long in dorsal aspect, the baso-

lateral sclerotization vertical; paraproct well sclerotized, with a single large, very

heavily sclerotized curved apical tooth; cercal setae fine, short, 3-6.

Zavortink: New World Aedes 13

PUPAE. Cephalothorax: Weakly to moderately pigmented, lighter ventrally. Hair

5-C moderately to strongly developed, short to moderately long, never reaching

more than 0.6 distance from its base to that of trumpet. Trumpet: Light golden

brown to dark brown in color. Broadening gradually from base to apex; trach-

eoid sculpturing relatively well developed in basal 0.11-0.21; reticulate sculptur-

ing strong. Abdomen: Weakly to moderately pigmented, lighter posteriorly. Ter-

gites II-VII without a strong transverse ridge anteriorly. Hair 1-II weakly to strong-

ly developed, usually 3-7b (1-14); hair 1-III-V subequally developed, weak to mod-

erately strong, usually with fewer [1-5 (1-8)], shorter and finer branches than 1-

II; hair 1-VI,VII weakly to moderately developed, subequal to 1-III-V or slightly

smaller, usually 1-4b (1-5). Hair 2-III-V mesad or laterad of hair 1 of corres-

ponding segment. Hair 5-IV,V 1.0-2.2 length of corresponding tergite; 5-VII mod-

erately long to long, usually single (single or double), weaker and shorter or strong-

er and longer than 4-VII. Hair 6-III-VI subequally developed, moderately long,

fine to moderately strong; 6-VII shorter and finer than 6-III-VI. Hair 8-VI,VII ven-

tral. Hair 9-III-VI subequally developed, short to relatively long, fine to stout,

caudad of the level of hair 6 of the corresponding segment (except possibly in

knabi); 9-VII near or moderately cephalad of caudolateral angle of segment, with

1 or 2 (1-4) moderately long to long primary branches; 9-VIII at or slightly mesad

of caudolateral angle of segment; with 3-5 (2-7) long; strong primary branches.

Hair 10-VI relatively close to 11-VI. Terminal Segments: Male genital lobe mod-

erately large, about 1.1-1.4 length of tergite VIII. Paddle: Length always greater

than width. Apex produced, pointed, rounded or subtruncate. Midrib conspicuous

to or nearly to apex; without wrinkled area near apex. Without long marginal

spicules. Hair 1-P short to moderately long, usually single (single or double).

LARVAE. Head: Lightly to moderately pigmented, lightened in ocular area,

darkened posteriorly. Labial plate distinctly narrowed anteriorly. Hair 1-C usu-

ally stout (fine in knabi). Hairs 4,6-C displaced caudad, not near labrum. Hair

4-C weakly to strongly developed, 2-24b; mesad or laterad of 1-C and cephalad

or caudad of level of 6-C. Hair 5-C single to multiple; more or less in line with

6-C. Hair 6-C single to multiple; distinctly laterad of 1-C. Hair 7-C usually 3-11b

(2-13). Hair 15-C usually short to moderately long and 2,3b (1-3) (long and 25-

35b in knabi). Mental plate with 9-12 (8-13) teeth on each side of median tooth.

Antenna: Shaft with or without small spicules. Hair 1-A moderately long, usually

single (1-3b). Thorax: Epidermis and fat body with or without pigment. Integu-

ment without spicules. Tubercles of hairs 5-7-P separate. Hair 1-M moderately

long to long, branched; 1-T usually moderately long, branched. Hair 4-P branched.

Hair 5-P usually branched (single in knabi); 5-M single. Hair 11-P usually much

shorter than 9-P, sometimes enlarged and up to 0.6 length of 9-P; hair 11-M,T

much shorter than 9-P. Hair 13-T moderately to strongly developed, usually 2-

7b (2-10); subequal in size to 14-M or slightly larger. Hair 14-M weakly to strong-

ly developed, usually 3-9b (2-11). Abdomen: Hair 1-I weakly to strongly devel-

oped, usually 3-8b (2-10); hair 1-IV,V strongly developed, usually 2-7b (1-8).

Hair 2-III-V laterad or mesad of hair 1 of corresponding segment; usually sin-

gle or double (1-4b) (3-6b and stellate on III and IV in chionotum). Hair 3-

VII weakly to moderately developed, usually 2-4b (1-5), short. Hair 5-II-IV mod-

erately to strongly developed, usually 2-7b (1-9). Hair 6-III-V usually double.

Hair 9-III-V usually single or double (1-3b). Hair 11-I weakly to strongly devel-

oped, usually 3-8b (2-11). Hair 12-I usually present (absent in zoosophus). Hair

13-I weakly to strongly developed, usually 2-6b (2-8); hair 13-IV,V strongly de-

14 Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

veloped, usually 2-8b (1-9); hair 13-VI usually with numerous [6-13 (3-18)] short

fine branches and usually cephalad of the level of 10-VI [with fewer (4) longer

and stronger branches and then sometimes distinctly cephalolaterad or laterad

of 10-VI in sandrae]. Segment VIII: Hairs 1 and 2 separated. Hair 1 moderate-

ly to strongly developed, usually 3-8b (1-9). Comb scales small to moderate in

size; fringed; relatively few (8-15) in 1 or more regular or irregular rows or num-

erous (16-58) in a patch. Siphon: Moderately to strongly pigmented. Short to

moderately long, index 1.7-5.0. Acus small to moderate in size. Pecten teeth

with numerous irregular basal denticles; 13-47 in a more or less straight even row.

Hair 1-S slightly distad of pecten; moderately to strongly developed, 1-5b. Anal

Segment: Saddle small to large; without spines on caudal margin; moderately to

strongly pigmented. Hair 1-X moderately to strongly developed, usually 2-7b (1-

9) (single in knabi). Hair 2-X usually 4-6b (3-9). Hair 3-X single. Ventral brush

(4-X) weakly to strongly developed; with 5 (hendersoni, brelandi), 6 (most species)

or 7 (zoosophus, knabi) pairs of hairs; all but most proximal 1 or 2 hairs on grid;

hair 4a-X usually long (shortened in sandrae), 3-6b; hairs 4b,4c-X usually 3,4b

(2-5) [regularly double (2,3b) in hendersoni, apparently 5-7b in knabi]. Anal gills

varied.

DISCUSSION. The subgenus Protomacleaya is characterized as follows: in 1 the

adults by the combination of (1) the narrowly separated to partially contiguous

eyes, (2) the usual presence of numerous erect scales and narrow curved scales

on the vertex of the head, (3) the ornamentation of the mesoscutum, which is

usually dark scaled with anterior lateral or lateral silver or white markings, (4)

the 3 distinct patches of scales on the sternopleuron, (5) the large patch of scales

on the paratergite and laterotergite, (6) the usually single large patch of scales

on the mesepimeron, (7) the completely scaled posterior pronotum with the

scales usually broad and flat, (8) the relatively stout proboscis, and (9) the us-

ually entirely dark scaled proboscis and palpus; in the male genitalia by (1) the

shape of the aedeagus, which is usually rounded and slightly expanded basally,

more or less parallel-sided distally, and truncate apically, (2) the frequent devel-

opment of at least a weak sternomesal sclerite and tuft, and the combination

of (3) the lobes of tergite IX, which bear 1-9 setae in a single series, (4) the

patch of few to very numerous weakly to moderately developed setae in the basal

tergomesal area of the sidepiece, (5) the absence of numerous long, strong se-

tae along the apical portion of the sternomesal surface of the sidepiece, (6) the

relatively short clasper without setae on the outer surface near the apex, and (7)

the relatively long spiniform; in the pupae by the combination of (1) the absence

of a transverse anterior ridge on tergites II-VII, (2) the usually strongly pigmented

trumpet, (3) the usually relatively strongly developed and long branches of hair

1-II, (4) the moderately long to long 5-IV,V, and (5) the usually fine 9-III-VI;

and, in the larvae by the combination of (1) the weakly to moderately developed,

short, 2-6b (1-7) hair 3-VII, (2) the moderately to strongly developed 5-II-IV,

(3) the usual presence of 12-I, (4) the comb scales, which are fringed and num-

erous enough to form a large patch in most species, and (5) the development

of the ventral brush, which arises from a grid and is moderately to strongly de-

veloped, with 6-8 pairs of hairs and 4b,4c-X 3-12b, in most species.

Protomacleaya, with more than 40 species, is the dominant subgenus of tree-

hole-breeding Aedes in the New World. Although the subgenus is quite diverse,

is defined mainly on negative characters, and many of the included species depart

from the “‘normal’’ in 1 or more characteristics of 1 or more stages, it is, I believe,

Zavortink: New World Aedes 15

natural because of the basically similar male genitalia of virtually all species. There

is no morphological evidence to indicate that Protomacleaya is related to any

other Middle American group of treehole-breeding Aedes, and I suspect that this

subgenus has been produced by radiation following the independent, relatively

recent invasion of this habitat by a ground-pool breeding form in Middle America.

To the north, in Mexico, the United States, and Canada, the distributions of

Protomacleaya and the Pulchritarsis Section of Ochlerotatus are basically com-

plementary; while I would usually interpret this as a sign of relationship, I do

not believe that can be the explanation in this case.

The species of Protomacleaya fall into 5 groups. Four of these, the monotypic

Knabi and Zoosophus Groups and the larger Kompi and Triseriatus Groups, are

treated in the present study; the fifth and largest group, Terrens, has been re-

vised by Schick (1970a,b). The 5 groups appear to be distinguishable only in the

adult and larval stages. Even the 4 small groups treated here can be separated

only with difficulty in the pupal stage and male genitalia.

Since the groups of Protomacleaya differ most conspicuously in adult orna-

mentation, particularly tarsal banding, their relationship was not detected by Ed-

wards (1932) or Knight and Marks (1952), and the species of Protomacleaya

known to these authors are scattered throughout several of the major groups (B,

F; and H) of ':Finiaya:”

The description of Protomacleaya is based on only the 4 groups treated here;

the diagnosis, though, does include the Terrens Group, which has also been added

to the keys to the adults and larvae.

The immatures of Protomacleaya are found primarily in treeholes and occa-

sionally in broken or cut bamboo internodes, artificial containers, rockholes, co-

conut shells and pitcher plants.

Protomacleaya extends from southern Canada to northern Argentina and south-

eastern Brazil; it is absent from the West Indies. The Triseriatus, Kompi and Ter-

rens Groups replace each other geographically; the Zoosophus Group occurs with

the Triseriatus Group in the southcentral United States and the Knabi Group is

found with the Terrens Group in southern Mexico.

KEYS TO GROUPS

ADULTS

iy, Integument of head and thorax yellow to tan; scales of head and meso-

scutum largely yellow to golden; claws of hindleg with an acute sub-

median tooth in both sexes(11.knabi) . . . . .. . . Knabi Group

Integument of head and thorax brown to Black: scales of head and meso-

scutum usually largely dark brown to black and white to silver-white,

sometimes partially yellowish, creamish-tan or golden brown (Zoosophus

Group); claws of hindleg simple in both sexes

2(1). Tarsal segments 1-5 of all legs entirely dark scaled . . . Triseriatus Group

Some tarsal segments marked with white .........-.. .3

3(2). Tarsal segments 1-3 or 4 of foreleg and midleg and tarsal segments 1-4 or

5 of hindleg with conspicuous basal white band (4. zoosophus) .

Zoosophus Group

16

4(3).

2(1).

64 8

3(2).

2(1).

Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

Tarsal segments 3-5 usually entirely dark scaled . . ....... .4

Hindtarsus with a conspicuous band over joint between segments 1 and 2

and/or hindfemur with a dark basal band. . . . . . .lerrens Group

Hindtarsus without a band over joint between sepments 1 and 2; hind-

femur without adark basalband. . ..... . . . .Kompi ‘Group

MALE GENITALIA

(Terrens Group not included)

Setae of basal tergomesal area of sidepiece arising from a differentially

sclerotized plaque (11. knabi). . . . . . . . . Knabi Group

Setae of basal tergomesal area of Sidepieds. not Aeitis from a differentially

Sects ere me er ee oe a en ae oe see ag

Aedeagus relatively short and broad (length 1.7-2.0 times maximum width)

and median sternomesal tuft of sidepiece weakly to moderately developed

. . . Kompi Group

ener itively jong anid sender (engi i : -2. 3 times maximum width)

or median sternomesal tuft of sidepiece strongly developed .... .3

Median sternomesal tuft of sidepiece very weakly developed and incon-

spicuous (4. zoosophus) . . . . . . . Loosophus Group

Median sternomesal tuft of sidépiecé nisaeiately to strongly developed and

conspicuous to very conspicuous. . .. . . . . . Triseriatus Group

PUPAE

(Terrens Group not included)

Hairs 4,8-C very strongly developed and long . . . . . . Kompi Group

Hairs 4 ,8-C weakly to strongly developed and short a, See ee or es

Hair 6-I,II strongly developed, as stout as or stouter than 3-I,II; hair 9-III-

VI stout and relatively long (4. zoosophus). . . . . Zoosophus Group

Hair 6-I,1] moderately developed, much finer than 3-I,II; hair 9-III-VI fine,

SET a PTV OT ue ae ee gas i ean vip te ae

Apex of paddle rounded or subtruncate . . . . . . Triseriatus Group

Apex of paddle pointed or produced (11. knabi).. . . . . » Knabi Group

LARVAE

Hair 15-C long, 25-35b; hair 1-C fine; mental plate with 1 or 2 lateral

teeth much enlarged (11. knabi) . . . . . . Knabi Group

Hair 15-C short to moderately long, 1-3b: Be 1-C stout; mental plate

without conspicuously eniarecdlaloral.f6eth yo oes ek eee se 2

Hair 4-C well developed, with 8-24 long branches, nearer to longitudinal mid-

line of head than to 6-C; comb scales relatively few, 10-13 (8-15), in

1 or 2 regular or irregular rows ee Pane ; . a

Zavortink: New World Aedes 17

Hair 4-C weakly developed, with 2-7 (2-11) short branches, usually nearer

to 6-C than to longitudinal midline of head; comb scales relatively num-

érous, 20-100 asually aw ar patel i> 6g cers dat a ater oo Od ae

3(2). Saddle of anal segment large, extending 0.65-0.80 distance down lateral

surface of segment, with a narrow lightened or transparent ventral mar-

ginal or submarginal area; ventral brush tie with 7 pairs of hairs; hair

12-I absent (4. zoosophus) . . . } . . . Zoosophus Group

Saddle of anal segment small to moderate in size, extending 0.40-0.65 dis-

tance down lateral surface of segment, without a narrow lightened or

transparent ventral marginal or submarginal area; ventral brush with 5

or 6 pairs of hairs; hair 12-I present. . . . . . . . Triseriatus Group

4(2). Ventral brush with 6 pairs of hairs and 4b,4c-X usually 3,4b (2-5)

. . Kompi Group

Vevtual brush with 5.9 pili of hairs: i witli éi pairs then 4b ,4c-X usually

ST Oba S- 23005 isis, ‘ : PeLURY Jot: Terrens4sroup

TRISERIATUS GROUP

FEMALES. Head: Integument brown to black. Erect scales usually unicolorous,

white to amber. Narrow curved scales usually unicolorous, white to silver-white

(dark brown in Florida triseriatus). Thorax: Integument brown to black. Acros-

tichal, anterior dorsocentral, humeral, lateral prescutal and posterior fossal bristles

numerous and well developed or much reduced in number and size or even mostly

absent except on anterior promontory. Mesoscutum with background of small dark

brown or black scales with varied pattern of larger white, silver-white, dingy silver-

white or even yellowish or tannish scales (Florida triseriatus), as follows: (1) short

to long broad patch on median anterior promontory usually developed, (2) nar-

row broken or complete acrostichal line sometimes developed, (3) narrow to broad

lateral prescutellar line usually developed, (4) narrow broken or complete poster-

ior outer dorsocentral line sometimes developed, and (5) broad to very broad

lateral marginal line extending from anterior promontory to wing root and mesad

along scutal suture usually developed. Midlobe of scutellum with scales dense,

mostly broad and flat, usually silver, sometimes dark. Ppn scales predominantly

broad and flat, usually white to silver-white; psp scales present or absent; ssp scales

usually present; pst and metameron normally bare. Legs: Tarsi entirely dark scaled

or with an inconspicuous pale longitudinal streak on posterior and/or ventral sur-

face of 1 or more basal segments. Claws of hindleg simple. Wing: Base of costa

with or without a small light patch. Base of vein R with or without a very small

light patch. Abdomen: Tergites II-VII usually dark scaled with basolateral light

patch. Basal sternites light scaled; distal dark scaled with basolateral light patch

or light basal band that is broader laterally.

MALES. Essentially as in females except for sexual characters. Head: Palpus

entirely dark scaled; with few to numerous long bristles from apex of segment

3 distad. Thorax: Mesoscutum usually more extensively light scaled than in fe-

males, rarely entirely light scaled. Legs: Larger claw of foreleg and midleg with

or without an acute basal external tooth; claws of hindleg simple. Wing: Remig-

ium (base of vein R) dark scaled or entirely or partially light scaled to about

level of crossvein h.

18 Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

MALE GENITALIA. Sidepiece: Usually weakly to moderately pigmented. Basal

tergomesal area slightly to conspicuously swollen; setae of basal tergomesal tuft

few to numerous, not arising from a differentially sclerotized plaque; median sterno-

mesal sclerite moderately to strongly developed; median sternomesal tuft mod-

erately to strongly developed, the setae sometimes strongly curved dorsad. Clasp-

ette: Filament evenly curved or angulate. Phallosome: Aedeagus relatively long

and slender, length usually 2.0-2.3 (1.8-2.4) greatest width.

PUPAE. Hairs largely moderately strong and moderately pigmented. Cephalo-

thorax: Hairs 4,8-C weakly to moderately developed, short; 8-C shorter and finer

than 9-C. Trumpet: Light golden brown to brown. Abdomen: Branches of hair

1-II weakly to moderately developed, short. Hair 6-I,II moderately developed, us-

ually much finer than 3-I,II. Hair 9-III-VI fine, short to relatively long. Paddle:

Apex rounded or subtruncate.

LARVAE. Head: Labrum more or less evenly rounded in dorsal aspect. Hair

1-C stout, arising on or near front edge of labrum. Hair 4-C well developed, with

numerous (8-24) long branches; closer to midline than to 6-C. Hair 5-C 1-3b.

Hair 6-C 1-5b. Hair 7-C 5-13b. Hair 15-C short to moderately long, 1-3b. Mental

plate distinctly triangular, lateral teeth not conspicuously enlarged. Antenna: Shaft

smooth or with a few spicules. Thorax: Pigment present or absent. Hairs 1,5-P

branched. Abdomen: Hair 12-I present. Segment VIII: Comb scales usually 10-

13 (9-15), in 1 or 2 irregular rows. Siphon: Index 2.2-3.5. Acus attached or de-

tached, sometimes removed from base of siphon. Pecten teeth short to long, mod-

erately to darkly pigmented; 13-30. Anal Segment: Saddle small to moderate in

size, extending 0.40-0.65 down lateral surface of segment; without a lightened

or transparent ventral marginal or submarginal area. Hair 1-X 1-9b. Ventral brush

with 5 or 6 pairs of hairs; hairs 4b,4c-X usually 2-4b (2-5). Gills sausage-shaped

or tapered; dorsal subequal to ventral or longer, 0.5-4.5 length of saddle.

DISCUSSION. The Triseriatus Group is characterized as follows: in the adults

by the dark tarsi; in the male genitalia by the combination of (1) the weak to

moderate pigmentation, (2) the absence of a differentially sclerotized plaque in

the basal tergomesal area, (3) the moderately to strongly developed sternomesal

tuft, and (4) the usually relatively long slender aedeagus; in the pupae by the com-

bination of (1) the short hairs 4,8-C, (2) the weak 6-I,II and 9-III-VI, and (3)

the rounded or subtruncate apex of the paddle; and in the larvae by the combina-

tion of (1) the well developed hair 4-C, (2) the usually 10-13 (8-15) comb scales

arranged in 1 or 2 irregular rows, and (3) the ventral brush, which has 5 or 6

pairs of hairs and 4b,4c-X 2-4b (2-5).

This group contains at least 3 species, triseriatus, hendersoni and brelandi. Aedes

triseriatus is conspicuously differentiated from the other 2 in the adult and larval

stages and male genitalia. Aedes hendersoni and brelandi differ in relatively few

characters of the adults, pupae and larvae.

The light scaling of the mesoscutum of species in this group is usually more

extensive in males than in females and is frequently less extensive in depauperate

individuals of both sexes. This must be considered when identifying specimens

on the basis of the mesoscutal pattern.

The immature stages of species in the Triseriatus Group are characteristically

found in rot cavities in trees, stumps and logs and in water-holding crotches of

trees. They have also been found in artificial containers, pitcher plants and rock-

holes. Females of all 3 species are known to bite humans.

The Triseriatus Group is restricted to North America, where it extends from

Zavortink: New World Aedes 19

southern Canada to northern Mexico. Two of the species, hendersoni and triser-

iatus, are widespread and sympatric; the third, brelandi, is presently known from

a limited area of southwestern Texas that is outside the range of both other

species. The distributions attributed to the species of this group are based solely

on the material that I have examined because, in most instances, it is impossible

to tell to which species published records of triseriatus refer.

2(1).

ET):

KEYS TO SPECIES

ADULTS

Bristles of anterior portion of mesoscutum relatively few and predominantly

weakly developed, the fossa not completely bordered by setae and rarely

more than | or 2 of the setae conspicuous; light scaling of fossa reduced,

usually restricted to a narrow or broad stripe along lateral and posterior

portions; base of costal vein of female completely dark scaled or with

only a few light scales; palpus of male with relatively few long bristles

from apex of segment 3 distad (fig. 2)... . . . . . 3. triseriatus

Bristles of anterior portion of mesoscutum relatively numerous and well

developed, the fossa more or less bordered on all sides by conspicuous

setae; light scaling of fossa usually extensive and extending mesad of

anterior dorsocentral bristles; base of costal vein of female usually with

small but conspicuous white patch; palpus of male with numerous long

bristles: tromcapexof seoment:.3 distad™ aig ks A ee Pe, Kees Se ee

Psp scale patch absent to small in female, usually absent in male; bristles

around fossa of female lightly to moderately pigmented (fig. 2)

. 1. hendersoni

Psp eeale patel farge i in female, Geually email. in ake prietics around fossa

of female more darkly pismented (f396.. 2 5) ar create, vat a ere ONebd

MALE GENITALIA

Tergal surface of sidepiece conspicuously constricted at level of claspette

filament; setae of mesal half of middle portion of tergal surface of side-

piece much reduced in size and number; median sternomesal tuft strongly

developed, many of the setae sharply curved cephalad and/or dorsad (figs.

Mos eet eee te . . . . IL. hendersoni; 2. brelandi

Tergal surface of sidepiece ‘not constricted: setae of mesal half of middle

portion of tergal surface of sidepiece not reduced in size or number;

median sternomesal tuft moderately developed, none or very few of the

setae sharply curved cephalad and/or dorsad (fig. 8) . . . 3. triseriatus

PUPAE

Trumpet light to medium golden brown in color (fig. 6). . . . 2. brelandi

Trompet medium 16 dark brown i COlOT 5 oe Ss sis cae

Hair 1-IV eer 4,5b BS ey hair 5-IV Ng yax, 1.6-2.2 hii of 3-III (fig. 3)

, ai . 1. hendersoni

20 Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

Hair 1-IV usually 2,3b (2- nae hair 5-1V soon 1.0-1.3 (1.0-2.0) length of

Sab Chie) oe! Leer ; oe . . . 3. triseriatus

LARVAE

1. Ventral brush (4-X) with 6 pairs of hairs; thorax and abdomen pigmented;

hair 1-X usually 4,5b (2-9); anal gills tapering distally, relatively short,

dorsal longer than ventral 1G as anes . . 3. triseriatus

Ventral brush with 5 pairs of hairs; thorax and ‘abdomen without pigment;

hair 1-X usually double (1-3b); anal gills sausage- ‘dain nasi nina

dorsal and ventral subequal in length . ; ae

2(1). Hairs 4b,4c-X usually double (2,3b) (fig. 4). . . . . . . . 1. hendersoni

Hairs 4b,4c-X usually 3,4b (2-5) (fig.7) . . . . . . . . . . 2. brelandi

1. Aedes (Protomacleaya) hendersoni Cockerell

Figs. 1-4

1918. Aedes triseriatus var. hendersoni Cockerell, 1918:199-200. TYPE: Lectotype female,

Box Elder Creek, 19 mi W Douglas, Wyoming, United States of America, 25 Aug 1917,

E. Schwabe and J. Henderson [USNM; selection of Stone and Knight, 1956:218]. Re-

duced to synonymy with triseriatus by Dyar (1919:39); resurrected to specific rank

by Breland (1960:601).

Aedes (Finlaya) hendersoni of Breland (1960:600-606); Stone (1961:40).

Aedes hendersoni of Hedeen (1963:349-350); Nielsen, Arnell and Linam (1967:72-74); Nielsen,

Linam, Arnell and Zavortink (1968:361-362); Truman and Craig (1968:1020-1025); Harmston

(1969:490-491); Loor and DeFoliart (1970:60,64). ,

Aedes (Finlaya) triseriatus in part of Dyar (1928:222); Edwards (1932:155); Matheson (1944:

187-188); Knight and Marks (1952:542,572); Carpenter and LaCasse (1955:255-257); Stone,

Knight and Starcke (1959:172); Stone (1961:41).

Aedes triseriatus in part of Howard, Dyar and Knab (1917:762-766); Dyar (1919:39); Rozeboom

(1942:34); Jenkins and Carpenter (1946:37-39); Breland (1949:93-100).

For additional references see Carpenter (1970:53-54).

FEMALE (fig. 2). Wing: 3.87 mm. Proboscis: 2.35 mm. Forefemur: 2.23 mm.

Abdomen: about 3.1 mm. Head: Decumbent scales of dorsal surface light. Thorax:

Most mesoscutal bristles lightly to moderately pigmented. Acrostichal bristles us-

ually restricted to anterior promontory; anterior dorsocentral, humeral, lateral pre-

scutal and posterior fossal bristles relatively numerous and conspicuous. Light

mesoscutal scales predominantly silver-white in western populations, frequently

more dingy in eastern populations. Narrow interrupted or complete acrostichal

and posterior outer dorsocentral light scale lines sometimes developed in western

populations; lateral prescutellar light scale line present; light scaling of fossa ex-

tensive, usually extending mesad of anterior dorsocentral bristles; light scales some-

times in anterior transverse band on mesoscutum in western populations. Mid-

lobe of scutellum with light scales. Pleural scale patches silver-white. Psp scales

absent or very few. Legs: Forecoxal scales usually all light. Posterior or ventral

surface of forefemur and midfemur with light scales restricted to basal 0.6-0.8

or extending to apex in a narrowed streak. Knee spot absent to moderate on

foreleg, small to moderate on midleg, moderate to large on hindleg. Tibiae and

Zavortink: New World Aedes 21

tarsi very frequently marked with pale scales on at least posterior surface. Claws

of foreleg and midleg rather abruptly and sharply curved, the tooth long. Wing:

Costa usually with small basal light patch. Haltere: Varying from entirely light

scaled to predominantly dark scaled. Abdomen: Tergites H-VII with large baso-

lateral white to silver-white patch that is usually rounded mesally in eastern pop-

ulations, usually cuneate mesally and sometimes joining mate in western populations.

MALE (fig. 2). Essentially as in female except for sexual characters. Head: Pal-

pus with numerous long bristles from apex of segment 3 distad. Thorax: Meso-

scutal bristles lighter than in female. Light scaling of mesoscutum frequently more

extensive than in female, rarely all scutal scales light. Psp scale patch usually

absent. Legs: Larger claw of foreleg and midleg with an acute basal external tooth.

Wing: Remigium (base of vein R) with light scales.

MALE GENITALIA (fig. 3). Sidepiece: Tergal sclerotization conspicuously broad-

ened distally, so that tergal surface appears constricted at level of claspette fila-

ment; basal tergomesal area usually conspicuously swollen and with numerous

elongate setae in a more or less oval to circular patch, the conspicuousness of

these setae enhanced by the reduction in size and number of the setae on the

mesal half of middle portion of tergal surface; median sternomesal sclerite strongly

developed, triangular, projecting far into mesal membrane; median sternomesal

tuft strongly developed, consisting of very numerous setae, many of which are

sharply curved cephalad and/or dorsad. Claspette: Filament angulate. Phallosome:

Aedeagus about 2.0-2.1 (1.8-2.2) times longer than greatest width.

PUPA (fig. 3). Abdomen: about 3.2 mm. Trumpet: 0.56 mm. Paddle: 0.90

mm. Trumpet: Medium to dark brown in color. Abdomen: Hair 1-II usually 4-6b

(3-14); hair 1-IIJ usually 4-6b (2-10); hair 1-IV usually 4,5b (2-8); hair 1-V_ us-

ually 3-5b (2-6). Hair 2-III usually mesad of 1-H. Hair 4-II usually laterad of

5-II. Hair 5-IV usually 1.6-2.2 length of 3-IIl.

LARVA (fig. 4). Head: 0.84 mm. Siphon: 0.85 mm. Anal Saddle: 0.26 mm.

Head: Hair 5-C usually single (1-3b). Hair 6-C usually 3,4b (3-5). Hair 8-C usu-

ally double (single or double). Thorax: Pigment absent, living larva white in color.

Alveolar plate of hairs 1-3-P usually absent. Abdomen: Hair 4-III,[V usually dou-

ble or bifid (1-3b or 3f). Siphon: Index usually 2.7-3.3 (2.5-3.5). Acus detached

and usually removed from base of siphon. Pecten teeth relatively short, usually

moderately pigmented; 18-26 (15-30) in number. Hair 1-S usually 2,3b (2-4). Anal

Segment: Saddle small to moderate in size, extending about 0.40-0.60 down lat-

eral surface of segment. Hair 1-X located near caudolateral angle of saddle or

less frequently along posterior margin of saddle near its caudolateral angle, its

alveolus usually at edge of saddle or in unsclerotized notch; usually double (1-

3b). Ventral brush with 5 pairs of hairs; hairs 4b,4c-X usually double (2,3b).

Gills sausage-shaped; dorsal and ventral subequal in development and long to very

long, usually 3.0-4.0 (2.4-4.4) length of saddle.

SYSTEMATICS. I am in full agreement with Breland (1960), who restored

hendersoni to full specific rank. This species is well differentiated from triseriatus

in the adult and larval stages and male genitalia and can be easily separated from

that species as follows: in the adult by (1) the numerous and well developed an-

terior dorsocentral, humeral, lateral prescutal and posterior fossal bristles, (2) the

more extensive light scaling of the fossal area, the light scales almost always ex-

tending mesad of the anterior dorsocentral bristles, (3) the claws of the fore-

leg and midleg, which are abruptly and sharply curved and have a long submed-

ian tooth in the female, and which have an external tooth at the base of the

22 Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

larger claw in the male, (4) the ornamentation of the wing, which has a small

but conspicuous patch of white scales at the base of the costa in the female

and the remigium (base of vein R) entirely or partially light scaled in the male,

and (5) the more numerous long bristles on the palpus of the male; in the male

genitalia most conspicuously by (1) the constricted tergal surface of the side-

piece, (2) the more numerous elongate setae of the basal tergomesal area of the

sidepiece and their arrangement in a more or less oval to circular patch, (3) the

reduction in size and number of the setae on the mesal half of the middle por-

tion of the tergal surface of the sidepiece, (4) the larger, triangular sternomesal

sclerite, (5) the strongly developed sternomesal tuft with many of the setae strong-

ly curved cephalad or dorsad, and (6) the more angular claspette filament; and

in the larva by (1) the absence of pigment in the thorax and abdomen, (2) the

acus, which is detached and removed from the base of the siphon, (3) the shorter

and usually more lightly pigmented pecten teeth, (4) the usually double (1-3b)

hair 1-X, (5) the ventral brush which has 5 pairs of hairs and 4b,4c-X usually

double (2,3b), and (6) the 4 subequally developed, long to very long sausage-

shaped anal gills. The pupae of hendersoni and triseriatus are not as reliably sep-

arated as the other stages of these species, but the pupa of hendersoni is usually

distinguished from that of triseriatus by (1) the more numerous branches in hair

1-IV, and (2) the longer 5-IV.

Aedes hendersoni is most closely related to brelandi, a species from the Chisos

Mountains of southwestern Texas, and is separated from it by relatively few char-

acters in each stage, as follows: in the adult by (1) the usual absence of acros-

tichal bristles on the disc of the mesoscutum, (2) the lighter color of the meso-

scutal bristles, (3) the more silver-white or dingy-white color of the light scales

on the thorax, and (4) the reduced number or even absence of postspiracular

scales; in the pupa by the darker trumpet; and in the larva by the more weakly

developed ventral brush in which hairs 4b,4c-X are usually double (2,3b). The

male genitalia of hendersoni and brelandi are apparently indistinguishable.

Adults of hendersoni from areas west of the range of triseriatus are generally

more extensively light scaled than those from within the range of that species.

Many from the west have light scales in a transverse band across the anterior

portion of the mesoscutum and in narrow acrostichal and outer posterior dorso-

central lines, and most from this area have. the basolateral light patches of the

abdominal tergites enlarged and distinctly cuneate. The basis for the east-west

variation has not been determined.

Since hendersoni is widespread in temperate North America and its larva and

male genitalia are among the most derived in this subgenus, it is undoubtedly

one of the most recent species of Protomacleaya.

BIONOMICS. The immature stages of hendersoni normally occur in treeholes;

according to Truman and Craig (1968:1021) they are found only in those tree-

holes with water of high organic content. The Cornell University collection con-

tains adults with the label “‘bred from pitcher plant” [Sarracenia purpurea Lin-

naeus]. Adults of both sexes of hendersoni have been taken at lights and resting

in buildings and females have been collected biting humans and visiting Ceanothus

flowers.

The immatures of hendersoni are frequently associated with 2 other species of

Protomacleaya, triseriatus and zoosophus, and sometimes with Ae. (O.) epactius and

Ae. (O.) sierrensis (Ludlow, 1905). It would be interesting to investigate the ecolog-

ical distinctions that permit the continued coexistence of these species of Proto-

\

Zavortink: New World Aedes 23

macleaya. This is particularly true in the case of hendersoni and triseriatus, fairly

closely related species that are geographically sympatric over a very broad area.

Either these species typically inhabit different types of forest, their immatures

characteristically occur in different sorts of treeholes, the larvae of each species

are competitively superior to those of the other in some particular kind of tree-

hole, or the larvae occupy different ecological niches within the same treehole.

The larva of hendersoni is neotenic, that is, it retains in the fourth instar char-

acteristics that are found in only the younger instars of less differentiated species

of the subgenus, such as the reduced extent of sclerotization and reduced ventral

brush. Since neoteny has obviously played a role in the differentiation of the

larvae of another pair of closely related arboricolous species from the eastern

United States, Orthopodomyia alba Baker, 1936 and O. signifera (Coquillett, 1896)

(Zavortink, 1968:50-51), it seems certain that this phenomenon bears upon the

question of competition between the members of these species pairs. The morph-

ology, physiology or behavior of these neotenic larvae must in some way adapt

them to a different type of treehole, make them competitively superior to their

less differentiated associates under some conditions, or change what they do in

a treehole. To postulate that competition between larvae of different species can

be reduced by neoteny forces one to question the extent of competition between

the different larval instars of a single species and the effect this might have on

the population structure of that species.

DISTRIBUTION (fig. 1). Aedes hendersoni is the most widespread treehole-

breeding mosquito in North America. I have seen specimens from all of the United

States of America (excluding Alaska and Hawaii) except Arizona, California, Flor-

ida, Nevada, North Dakota, Rhode Island, Vermont, Washington and Wisconsin.

The species has been reported from Wisconsin by Loor and DeFoliart (1970:60-

64), undoubtedly occurs in North Dakota, Rhode Island, Vermont and Washington,

and very probably occurs in northern Florida, which leaves only Arizona, Cali-

fornia and Nevada outside its range. I have seen specimens from the Canadian Prov-

ince of British Columbia and the species undoubtedly occurs throughout south-

ern Canada. It may also occur in northeastern Mexico. Material examined: 710

specimens; 169 males, 312 females, 174 larvae, 55 pupae; 23 individual rearings

(20 larval, 3 incomplete).

CANADA. British Columbia: Sidar, Kootenay Lake, 1 Aug 1968, L.T. Nielsen (N-25-68),

5 M, 1M gen, 2 F [UTAH].

UNITED STATES. Alabama: Wilson Dam, Colbert Co., July 1954, R.X. Schick, 1 L [UCLA].

Arkansas: Brinkley, 18 May 1923, H.G. Dyar, 1 F [USNM]. Camp Pike, 17 May 1918, R.W.

Gies, 1 F [USNM]. Scott, 1-10 June 1909, J.K. Thibault, 1 F [USNM]. Stone Co., 5 Aug

1915, 1 F [USNM]. Colorado: Aurora, 6 Sept 1944, L.E. Perry, 1 F [UTAH]. Boulder, 8

km NE, 8 July-15 Aug 1958, J.R. Hilliard, 3 M, 3 M gen, 8 F, 6 L [USNM]. Greeley, 4 July 1962,

F.C. Harmston, 1 M [UTAH]. Kuner, 5 June 1959, F.C. Harmston, 17 M, 2 M gen, 12 F

[CDC], 8 M, 14 F [USNM]. Platteville, 1 June 1962, F.C. Harmston, 8 M, 4 M gen, 2 F

[UTAH]. Pueblo, 5 Sept 1944, L.E. Perry, 1 F [UTAH]. Vineland, 30 May 1967, M. Manson,

5 M, 1 F [UTAH]. Connecticut: Redding, 4 July 1928-July 1930, A.L. Melander, 3 F [USNM].

Delaware: Newark, 6 May 1936, D. MacCreary, 1 M, 1 M gen [USNM]. District of Columbia:

Site not specified, 22 June 1906-25 June 1915, F. Knab, 2 F [USNM]; 9 May 1918, H.G.

Dyar, 1 M [USNM]; 14 May 1918, H. Morrison, 1 F [USNM]; 26 Sept 1921, A.N. Caudell,

1 F [USNM]; 23 May 1924, R.C. Shannon, 1 F [USNM]. Georgia: Atlanta, Fort McPherson,

1943, S.J. Carpenter, 1 M [UTAH]. Atlanta, 2 June 1948, H.D. Pratt, 2 F [CDC], 1 F [UTAH];

16 Sept 1951, G. Heid, 1 F [UCLA]; 25 Mar 1953, W.D. Sudia and R.H. Gogel, 4 M, 1 M

gen [CDC]. Chamblee, Apr 1951, H.D. Pratt, 1 L [CDC]. College Park, 28 Apr 1948, L.T.

Foote, 1 M, 1 M gen [CDC]. Worth Co., 31 Mar 1951, J.K. Wall, 1 L [CDC]. Idaho: Nampa,

24 Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

27 June 1945, F.C. Harmston, 1 F [UTAH]. Jilinois: Glencoe, July 1923, G.W. Edwards, 1 F

[USNM]. Oakwood, Camp Drake, 28 July 1941, G.T. Riegel, 1 F [UTAH]. White Heath, 30

June 1938, Dirks, 1 M gen [CU]. Indiana: Lafayette, July 1928, 1 M [USNM]. Jowa: Ames, 21

July 1919, F.C. Bishopp, 1 F [USNM]. Kansas: Manhattan, 10 Oct 1934, C.W. Sabrosky, 1 F

[USNM]. Kentucky: Corbin, 24 Aug 1904, H.S. Barber, 3 F [USNM]. Hickman, J.A. Pearson, 1

F [USNM]. Louisiana: Baton Rouge, 22 Apr-24 May 1903, 3 F [USNM]. Maine: Kittery Point,

R. Thaxter, 1 F [USNM]. Maryland: Baltimore, 30 July 1919, 1 F [UCLA]. Hyattsville, 14

Aug 1907, 1 F [USNM]. Plummer’s Island, July 1907-Mar 1908, R.P. Currie, 2 F [USNM];

July 1924, R.C. Shannon, 2 F [USNM]; 7 Sept, 1 F [USNM]. River View, Aug 1905, T.

Pergande, 3 M, 1 M gen, 1 F [USNM]. Silver Spring, 20 July 1940, F.C. Bishopp, 1 M [USNM].

Takoma Park, 1 F [USNM]. Massachusetts: Framingham, 16 Sept 1937, C.A. Frost, 1 F [USNM].

West Springfield, 26 July-13 Sept 1903, F. Knab, 2 F [USNM]. Michigan: East Lansing, 21

June 1942, C. Sabrosky, 1 M [USNM]. Minnesota: Detroit Lakes, 8,9 Sept 1940, W.A. Hoffman, 1

F [USNM]. Mississippi: Agricultural College, 4 May-12 July 1905, G.W. Herrick, 2 F [USNM].

Belzoni, 4 Aug, H.S. Barber, 1 F [USNM]. Missouri: Atherton, 12 June-July 1915, C.F. Adams, 1

M, 1 F [USNM]. Gumbo, 4 July 1941, W.M. Gordon, 1 F [CU]. Henson, 14 June 1918, 1 F

[USNM]. Jefferson Bks’, 24 May 1927, E.C. Bingham, 1 F [USNM]. Joplin, 7 July 1942, A.B.

Gurney, 1 F [USNM]. Kansas City, July 1951, 2 F [CDC]. Kirkwood, 13 June 1952, F.C.

Harmston, 2 M [UTAH]. Neosho, Camp Crowder, 26,28 June 1942, A.B. Gurney, 1 M, 1 F

[USNM]. St. Louis Co., 4 June-17 July 1940, W.M. Gordon, 1 M, 1 M gen, 2 F [CU]. Webster

Groves, Satterthwait, 1 F [USNM]. Montana: Drummond (3.2 km S), 16 Aug 1967, L.T. Nielsen,

J.H. Linam and J.H. Arnell (N-37-67), 1 M, 1 M gen, 1 F [UTAH]. Forsyth, 16 Aug 1964,

L.T. Nielsen and J.H. Linam (N-34-64), 7 M, 4 F, 3 P, 3 L [UTAH]. Glendive, 18 June 1922,

H.G. Dyar, 2 F [USNM]. Hall (9.7 km S), 16 Aug 1967, L.T. Nielsen, J.H. Linam and J.H.

Arnell (N-36-67), 4 F [UTAH]. Hamilton, 5 Aug 1965, J.H. Arnell (HA-18-65), 2 M, 4 F, 3 P, 3

L; same data (HA-19-65), 8 F [UTAH]. Jordon, 16 Aug 1964, L.T. Nielsen and J.H. Linam

(N-32-64), 1 F, 2 L [UTAH]. Landusky (32 km S), James Kipp State Park, 15 Aug 1964, L.T.

Nielsen and J.H. Linam (N-30-64), 11 M, 1 M gen, 38 F [UTAH]. Lolo, 5 Aug 1965, J.H.

Arnell (HA-20-65), 7 L [UTAH]. Miles City, 16 Aug 1964, L.T. Nielsen and J.H. Linam (N-33-

64), 3 F, 3 P [UTAH]. Missoula, 6 July 1917, H.G. Dyar, 1 F [USNM]. Mosby, 16 Aug 1964,

L.T. Nielsen and J.H. Linam (N-31-64), 3 M, 5 F, 20 L [UTAH]. Silver Star, 15 Aug 1967, L.T.

Nielsen, J.H. Linam and J.H. Arnell (N-34-67), 1 M, 4 F [UTAH]. Nebraska: Albion, 10 Aug

1963, 8 L [UTAH]. Niobrara, 24 Aug 1943, W.C. Reeves and P. Galindo, 1 F [USNM]. Norfolk,

5 Aug 1943, W.C. Reeves and P. Galindo, 2 M, 1 M gen [USNM]. New Hampshire: Center

Harbor, 24 June, H.G. Dyar, 1 F [USNM]. Durham, 8 Aug, H.G. Dyar, 1 M, 1 M gen [USNM].

White Mountains, Morrison, 1 F [USNM]. New Jersey: Ridgewood, 28 June 1911, M.D. Leonard,

1 F [CU]. Union Co., 1938, 1 F [UTAH]. New Mexico: Puerto de Luna, 17 Mar 1968, L.T.

Nielsen, J.H. Linam and J.H. Arnell (HA-28-68), 4 M, 1 M gen, 1 F [UTAH]. New York:

Cayuta Lake, June 1943, R. Matheson, 1 M, 1 M gen, 5 F [CU]. Fort Niagara, 27 Sept 1944,

1 F [CDC]. Franklin Co., Goldsmith, Aug 1955, J.N. Belkin, 1 F [UCLA]. Honeoye Falls, 7

Aug 1915, M.D. Leonard, 1 F [USNM]. Ithaca, Aug 1901, 1 F [CU]; 6 July 1930-4 May 1942, 6

M, 4M gen, 12 F, 3 L [CU] ; 19 Sept 1941-Apr 1942, J.N. Belkin, 1 M, 1 M gen, 2 F,3 P [UCLA].

Lake Mahopac, 13 July 1908, 1 F [CU]. McLean Bog, Aug 1942, H.T. Dalmat, 1 F [CU].

McLean Reservoir, The Shack, 7 July 1924, 1 F [CU]. Rome, 8 Oct 1944, 1 F [CDC]. Saranac,

30 Aug 1955, J.N. Belkin, 1 F [UCLA]. Suffolk Co., July-Aug 1928, 1 F [CU]. North Carolina:

Charlotte, 30 May 1916, H.P. Barret, 1 M, 1 F [USNM]. Henderson Co., 7 Oct 1901, J.L. Coker,

Jr., 1 F [USNM]. Lumberton, 13 May 1920, H.P. Barret, 1 F [USNM]. Old Fort, 28 Sept 1933,

CCC Survey, 9 F [USNM]. Ohio: Ravenna, 8 June 1970, M.J. Zavortink (UCLA 571), 7 lpM

(571-24,29,32,33,35,38,40), 6 IlpF (571-21,26,28,30,34,37), 3 Ip (571-25,36,43), 2 M gen, 1 F, 1

P, 5 L [UCLA]. Oklahoma: Flint, 19 June 1937, Standish-Kaiser, 1 F [CU]. Gore, 20 July

1937, Standish-Kaiser, 1 F [CU]. Page, 23 June 1937, Standish-Kaiser, 1 F [CU]. Wister, 2

July 1904, H.S. Barber, 1 F [USNM]. Woodward (9.7 km E), Boiling Springs State Park, 1 Sept

1968, L.T. Nielsen (N-32-68), 7 M, 1 M gen, 8 F, 9 P, 50 L [UTAH]. Oregon: Pendleton (3.2

km E), 18 Aug 1967, L.T. Nielsen, J.H. Arnell and J.H. Linam (N-41-67), 1 M, 1 M gen [UTAH].

Zavortink: New World Aedes 25

Pennsylvania: Allentown, 22 July 1945, R.C. Barnes, 1 F [CDC]. Philadelphia, University of

Pennsylvania, 28 July 1921, 1 F [USNM]. South Carolina: Richland Co., 10 Oct 1944, 1 F

[CDC]. South Dakota: Belle Fourche, 14 July 1966, J.H. Linam, J.H. Arnell and L.T. Nielsen, 1

F [UTAH]. Hot Springs (9.7 km SE), 14 July 1966, J.H. Arnell, JH. Linam and L.T. Nielsen

(HA-34-66), 8 M, 9 F, 4 P, 15 L [UTAH]. Tennessee: Reelfoot Lake, 1 Sept 1941, M. Gordon, 1

F [CU]. Rives, 27 July, H.S. Barber, 1 F [USNM]. Texas: Austin, 1 Sept 1957, Zilker, 2 L

[UTAH]. Huntington, 24 Apr 1934, CCC Survey, 1 F [USNM]. Leon Springs, Camp Stanley, 24

May 1918, D.L. VanDine, 1 F [USNM]. Matagorda Co., 13 Aug 1943, 1 F [CDC]. Utah: Weber

Co., Weber Canyon, 13 May 1970, L.T. Nielsen et al. (HA-7-70), 1 lpM (7), 6 lpF (1,2,4,5,6,8), 6

M, 1 M gen, 1 F, 2 L [UTAH]. Virginia: Fairfax Co., Dead Run, 22 June 1915, R.C. Shannon, 1

F [USNM]. Falls Church, Holmes Run, 4 June-1 July 1961, W.W. Wirth, 1 M, 1 F [USNM].

Falls Church, 4 July 1913, F. Knab, 1 M, 1 M gen [USNM]. Glencarlyn, 26 June 1910, F. Knab,

1 F [USNM]. Monterey, 20 Aug 1934, S.S. Cook, 1 F [USNM]. West Virginia: Rock Gap,

Cacapon State Park, 1-5 Sept 1955, C.W. Sabrosky, 2 F [USNM]. Wyoming: Careyhurst, Boxelder

Creek, 15 July 1966, L.T. Nielsen, J.H. Arnell and J.H. Linam (HA-41-66), 5 M, 1 M gen, 4 F,4 L

[UTAH] . Casper, 15 July 1966, L.T. Nielsen, J.H. Arnell and J.H. Linam (HA-42-66), 1 M, 11 F

[UTAH]. Devils Tower, 13 July 1966, J.H. Arnell (HA-31-66), 6 F [UTAH]. Lander, 12 July

1966, J.H. Arnell (HA-25-66), 1 M, 2 F [UTAH]. Lingle (3.2 km W), 15 July 1966, L.T. Nielsen,

J.H. Arnell and J.H. Linam (HA-38-66), 5 M, 7 F [UTAH]. Newcastle (47 km S), 14 July 1966,

L.T. Nielsen, J.H. Linam and J.H. Arnell (HA-35-66), 5 M, 1 M gen, 4 F, 2 P, 6 L [UTAH].

Niobrara Co., Old Woman Creek at Hwy. 85, 14 July 1966, L.T. Nielsen, J.H. Arnell and J.H.

Linam (HA-36-66), 2 M, 2 M gen, 3 F, 4 P, 6 L [UTAH]. Orin, 30 July 1961, L.T. Nielsen, 1

F [UTAH]. Orin Junction, 15 July 1966, L.T. Nielsen, J.H. Arnell and J.H. Linam (HA-40-66),

1 M [UTAH]. Riverton, 12 July 1966, L.T. Nielsen, JH. Arnell and J.H. Linam (HA-26-66),

4 M, 1 F; same data (HA-27-66), 4 M, 4 F, 6 L [UTAH]. State Unknown: Wilsor, 7 July 1931,

1 F [CU].

LOCALITY UNKNOWN. 1 M gen (550310-21) [UCLA].

2. Aedes (Protomacleaya) brelandi Zavortink, n.sp.

Figs. 1,2,5-7

TYPES: Holotype male with associated larval and pupal skins and genitalia (UCLA 611-11),

Chisos Mountains, Big Bend National Park, Brewster Co., Texas, United States of America, ele-

vation above 1770 m, larva from rothole in living evergreen oak tree, 1 Sept 1969, T.J. Zavortink

[USNM]. Allotype female with associated larval and pupal skins (UCLA 611-15), same data as

holotype [USNM]. Paratypes: 2 IpM (611-12,13), 2 IpF (611-14,18), 1 lp (611-17), 1 M gen,

1 L, same data as holotype (UCLA 611); 5 IpF (609-10-13,80), 2 pM (609-100,102), 5 pF

(609-101,103-105,108), 1 P, 4 L, same data as holotype (UCLA 609); 2 IpF (610-10,11), same

data as holotype (UCLA 610); 1 pM (612-102), 2 F, 2 P, 1 L, same data as holotype (UCLA

612); 4 IpM (613-15,20,22,27), 15 lpF (613-10-12,16-18,23-26,28-32), 2 lp (613-19,21), 4 pM

(613-100,102-104), 50 M, 4 M gen, 26 F, 134 P, 20 L, same data as holotype (UCLA 613);

3 IpF (614-10-12), same data as holotype (UCLA 614); 23 L, same data as holotype except

collected on 31 Aug 1969 by T.J. Zavortink and J.A. Bergland (UCLA 602); 2 IpM (603-29,

30), 3 IpF (603-32,36,90), 3 L, same data as holotype except collected on 31 Aug 1969 by

T.J. Zavortink and J.A. Bergland (UCLA 603); 1 L, same data as holotype except collected

on 31 Aug 1969 by T.J. Zavortink and J.A. Bergland (UCLA 604); 1 F, same data as holo-

type except collected biting human on 31 Aug 1969 by T.J. Zavortink and J.A. Bergland (UCLA

605); 1 F, same data as holotype except collected biting human on 31 Aug 1969 by T.J. Zavortink

and J.A. Bergland, and 2 IpM (606-10,11), 1 lp (602-12), 1 P, 3 L reared from eggs laid by F

(UCLA 606) [UCLA, ISET, BM]; 1 F, Chisos Basin Campground, Chisos Mountains, Big Bend Na-

tional Park, 10 Sept 1968, L.T. Nielsen (A-11-68) [UTAH]. This species is dedicated to Osmond

P. Breland in recognition of his contributions to the knowledge of the Culicidae.

26 Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

FEMALE (figs. 2,5). Wing: 3.58 mm. Proboscis: 2.17 mm. Forefemur: 2.05

mm. Abdomen: about 3.3 mm. Head: Decumbent scales of dorsal surface light.

Thorax: Most mesoscutal bristles darkly pigmented. Acrostichal bristles frequently

in a more or less complete row; anterior dorsocentral, humeral, lateral prescutal

and posterior fossal bristles numerous and conspicuous. Light mesoscutal scales

usually whitish. Narrow interrupted to moderately broad complete acrostichal and

posterior outer dorsocentral light scale lines frequently developed; lateral prescu-

tellar light scale line developed; light scaling of fossa extensive, extending mesad

of anterior dorsocentral bristles; light scales usually in anterior transverse band

on mesoscutum. Scales of midlobe of scutellum light. Pleural scales white. Psp

with well developed scale patch. Legs: Forecoxal scales all light. Posterior or ven-

tral surface of forefemur and midfemur with light scales restricted to basal 0.55-

0.70. Knee spot absent to small on foreleg, small to moderate on midleg, mod-

erate on hindleg. Tibiae and tarsi entirely dark scaled or rarely marked with broken

pale streak or pale speckles on posterior surface. Claws of foreleg and midleg

rather abruptly and sharply curved, the tooth long. Wing: Costa with small basal

light patch. Haltere: Light scaled. Abdomen: Tergites II-VII with large basolateral

cream-white patch that is usually rounded mesally.

MALE (figs. 2,5). Generally as in female except for sexual characters. Head:

Palpus with numerous long bristles from apex of segment 3 distad. Thorax: Meso-

scutal bristles frequently lightly pigmented. Light scaling of mesoscutum frequently

more extensive than in female, rarely all scutal scales light. Psp scale patch usually

present, small. Legs: Larger claw of foreleg and midleg with an acute basal ex-

ternal tooth. Wing: Remigium (base of vein R) with light scales.

MALE GENITALIA (fig. 6). Apparently indistinguishable from hendersoni.

PUPA (fig. 6). Abdomen: about 3.4 mm. Trumpet: 0.59 mm. Paddle: 0.91

mm. 7rumpet: Light to medium golden brown in color. Abdomen: Hair 1-II 4-

7b; hair 1-I[I-V usually 3-5b (2-6). Hair 2-III usually mesad of 1-III. Hair 4-Il

usually laterad of 5-II. Hair 5-[V usually at least 2.0 (1.9-2.7) length of 3-III.

LARVA (fig. 7). Head: 0.86 mm. Siphon: 0.78 mm. Anal Saddle: 0.25 mm.

Head: Hair 5-C usually double (1-3b). Hair 6-C usually 3,4b (2-6). Hair 8-C us-

ually double (single or double). Thorax: Pigment absent, living larva white in

color. Alveolar plate of hairs 1-3-P absent or poorly developed. Abdomen: Hair

4-IIT,1V usually 2-4b or 4f. Siphon: Index usually 2.5-3.2 (2.2-3.5). Acus detach-

ed and removed from base of siphon. Pecten teeth relatively short, usually mod-

erately pigmented; 18-24 (14-25) in number. Hair 1-S usually 3,4b (3-5). Anal

Segment: Saddle small to moderate in size, extending about 0.50-0.60 down lat-

eral surface of segment. Hair 1-X located along posterior margin of saddle near

its caudolateral angle or less frequently at caudolateral angle, its alveolus usually

at edge of saddle or in unsclerotized notch; usually double (1-3b). Ventral brush

with 5 pairs of hairs; hairs 4b,4c-X usually 3,4b (2-5). Gills sausage-shaped; dorsal

and ventral subequal in length and long to very long, usually 2.7-4.0 (2.3-4.5)

length of saddle.

SYSTEMATICS. Aedes brelandi is most closely related to hendersoni. It dif-

fers from that species as follows: in the adult by (1) the frequent presence of

acrostichal bristles on the disc of the mesoscutum, (2) the darker color of the

mesoscutal bristles, (3) the whiter color of the light scales of the thorax, and

(4) the greater number of postspiracular scales; in the pupa by the lighter color

of the trumpet; and in the larva by the more strongly developed ventral brush,

in which hairs 4b,4c-X are usually 3,4b (2-5). The male genitalia of brelandi and

Zavortink: New World Aedes 27

hendersoni are apparently indistinguishable.

The extent of light scaling on the mesoscutum of brelandi appears to vary

directly with size of the adults; the light acrostichal and outer posterior dorso-

central scale lines, so conspicuous on robust specimens, may be completely ab-

sent on smaller individuals.

BIONOMICS. The immature stages of brelandi were recovered from rotholes

in oak trees (Quercus spp.) growing at elevations above 1770 meters in the Chisos

Mountains. They were associated with the other arboricolous species Aedes (Och-

lerotatus) muelleri, Anopheles (An.) judithae Zavortink, 1969 and Orthopodomyia

kummi Edwards, 1939. Females of brelandi were troublesome biters in the vicin-

ity of these treeholes.

Rainfall in the Chisos Mountains was less than normal during the summer of

1969 and many treeholes that typically would have contained water in that sea-

son were dry. Many brelandi and muelleri were obtained as first instar larvae by

filling dry treeholes with drinking water and removing it 2-5 hours later.

DISTRIBUTION (fig. 1). Aedes brelandi is at present known from only the

Chisos Mountains in southwestern Texas. It was not recovered from numerous

collections made in the Davis Mountains to the northwest, but may occur to the

south at appropriate elevations in the Sierra Madre Oriental of Mexico. Such a

thesis is supported by the fact that it was collected with other treehole-breeding

species which occur farther south in the highlands of northern Mexico. Material

examined: 433 specimens; 68 males, 67 females, 102 larvae, 196 pupae; 58 in-

dividual rearings (42 larval, 12 pupal, 4 incomplete).

UNITED STATES. Texas: Chisos Mts., Big Bend National Park, Brewster Co., type series,

cited above.

3. Aedes (Protomacleaya) triseriatus (Say)

Figs. 1,2,8,9

1823. Culex triseriatus Say, 1823:12. TYPE: Female, Pennsylvania, United States of America

[nonexistent]. A neotype must be designated for triseriatus in order to preserve the

current taxonomic interpretation of this species and insure nomenclatorial stability. This

has not been done in the present paper because reared material from Pennsylvania is

not available.

1905. Finlaya ? nigra Ludlow, 1905:387-388. TYPE: Holotype female, Rock Island Arsenal,

Illinois, United States of America, 3 Aug 1905, G.G. Craig [USNM]. Synonymy with

triseriatus by Coquillett (1906c:21). This synonymy confirmed through courtesy of

W.W. Wirth.

Aedes (Protomacleaya) triseriata of Theobald (1907:254-257).

Aedes (Finlaya) triseriatus of Dyar (1928:222, in part); Edwards (1932:155, in part); Matheson

(1944:187-188, in part); Darsie (1951:14); Knight and Marks (1952:542,572, in part); Car-

penter and LaCasse (1955:255-257, in part); Stone, Knight and Starcke (1959:172, in part);

Breland (1960:600-606).

Aedes triseriatus of Dyar and Knab (1906:195-196); Howard, Dyar and Knab (1917:762-766,

in part); Cockerell (1918:200); Dyar (1919:39, in part); Bennett, Baker and Sellards (1939:

101-105); Davis (1940:49); Jakmauh (1940:17); Jenkins and Carpenter (1946:37-39, in part);

Breland (1949:93-100, in part); Chamberlain, Sikes, Nelson and Sudia (1954:280-283); Davis,

Hogge, Corristan and Ferrell (1966:227-230); Kissling and Chamberlain (1967:68); Truman

and Craig (1968:1020-1025); Harmston (1969:490-491); Loor and DeFoliart (1970:60-64);

Masterson, Stegmiller, Parsons, Croft and Spencer (1971:93-94,95); Messersmith (1971:7);

28 Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

Sudia, Newhouse, Calisher and Chamberlain (1971:584,590,595-596); Sudia, Newhouse and

Henderson (1971:209); Thompson, Anslow, Hanson and DeFoliart (1972:94-95).

Aedes (Gualteria) triseriatus of Dyar (1918:73,79); Vargas (1950:62); Vargas and Downs (1950:

171)

Ochlerotatus triseriatus of Coquillett (1906c:19,21).

For additional references see Howard, Dyar and Knab (1917:762-763), Carpenter and LaCasse

(1955:256-257), and Carpenter (1968:83; 1970:54).

FEMALE (fig. 2). Wing: 4.10 mm. Proboscis: 2.47 mm. Forefemur: 2.44 mm.

Abdomen: about 3.0 mm. Head: Decumbent scales of dorsal surface usually all

light in typical populations, dark in Florida population. Thorax: Mesoscutal bris-

tles lightly to darkly pigmented. Acrostichal bristles usually restricted to anter-

ior promontory; anterior dorsocentral, humeral, lateral prescutal and posterior fos-

sal bristles relatively few and predominantly inconspicuous in typical populations,

largely absent in populations from the southeastern United States. Light meso-

scutal scales silver-white in typical populations, tinged with yellow or tan in Flor-

ida population. Light acrostichal and posterior outer dorsocentral scale lines never

developed; lateral prescutellar light scale line present in typical populations, some-

times not developed in Florida population; in typical populations light scaling

of fossa usually restricted to a narrow or broad stripe along lateral and posterior

portions, but rarely extending mesad to dorsocentral area; light scaling usually

restricted to a small patch dorsocephalad of ppn or a very narrow lateral line

in Florida population; light scales rarely in complete anterior transverse band on

mesoscutum. Scales of midscutellar lobe usually all light in typical populations,

dark in Florida population. All pleural scale patches silver-white in typical popu-

lations, dorsal patches tinged with yellow or tan in Florida population. Psp scales

absent or very few. Legs: Forecoxal scales usually all light in typical populations,

partially dark in Florida population. Posterior or ventral surface of forefemur and

midfemur with light scales usually extending to apex in a broad streak in typical

populations, but with light scales usually restricted to basal 0.5-0.6 in Florida

population. Knee spot absent to small on foreleg, small to moderate on midleg,

and usually moderate on hindleg in typical populations, absent to small on fore-

leg and midleg and small on hindleg in Florida population. Tibiae and tarsi en-

tirely dark scaled or sometimes marked with pale streak on posterior or ventral

surface. Claws of foreleg and midleg usually evenly curved, the tooth usually short.

Wing: Costa usually entirely dark scaled at base, sometimes with a few light scales.

Haltere: Usually entirely or largely light scaled in typical populations, dark scaled

in Florida population. Abdomen: Tergites II-VII with basolateral white to silver-

white patch that is rounded mesally, the patch usually large in typical popula-

tions, usually small in Florida population.

MALE (fig. 2). Essentially as in female except for sexual characters. Head:

Palpus with relatively few long bristles from apex of segment 3 distad. Thorax:

Mesoscutal bristles frequently lighter than in female. Light scaling of mesoscutum

more extensive than in female, that of fossa frequently extending mesad to dorso-

central area. Psp scales absent. Legs: Larger claw of foreleg and midleg without

basal external tooth. Wing: Remigium (base of vein R) entirely dark scaled.

MALE GENITALIA (fig. 8). Sidepiece: Tergal sclerotization progressively nar-

rowing distally, the tergal surface not appearing constricted; basal tergomesal area

usually only slightly swollen and with relatively few elongate setae in a narrow

strip, the conspicuousness of these setae not enhanced by a reduction in size or

number of the setae on the mesal half of middle portion of tergal surface; med-

Zavortink: New World Aedes 29

ian sternomesal sclerite moderately developed, rounded, extending only moderate-

ly far into mesal membrane; median sternomesal tuft moderately developed, con-

sisting of moderately numerous setae, none or very few of which are sharply

curved cephalad and/or dorsad. Claspette: Filament more or less evenly curved.

Phallosome: Aedeagus about 2.2-2.3 (2.1-2.4) times longer than greatest width.

PUPA (fig. 8). Abdomen: about 3.7 mm. Trumpet: 0.51 mm. Paddle: 0.84

mm. Trumpet: Medium to dark brown in color. Abdomen: Hair 1-II usually 2-5b

(1-14); hair 1-III usually 2-4b (2-6); hair 1-IV usually 2,3b (1-4); hair 1-V 2,3b

(1-3). Hair 2-III usually laterad of 1-III. Hair 4-II usually mesad of 5-II. Hair

5-IV usually 1.0-1.3 (1.0-2.0) length of 3-III.

LARVA (fig. 9). Head: 0.92 mm. Siphon: 0.81 mm. Anal Saddle: 0.31 mm.

Head: Hair 5C single. Hair 6-C usually 2,3b (1-4). Hair 8-C usually single (single

or double). Thorax: Pigment present, living larva dark in color. Alveolar plate of

hairs 1-3-P usually well developed. Abdomen: Hair 4-III,IV usually single (single,

double or bifid). Siphon: Index usually 2.4-2.8 (2.3-3.0). Acus attached or de-

tached but near base of siphon. Pecten teeth relatively long, usually darkly pig-

mented; 16-25 (13-30) in number. Hair 1-S usually double (single or double).

Anal Segment: Saddle usually moderate in size, extending 0.50-0.65 down lateral

surface of segment. Hair 1-X usually located along posterior margin of saddle

near its caudolateral angle, its alveolus usually on saddle and removed from edge

of sclerotization by 2 or 3 alveolar diameters; usually 4,5b (3-9) in typical pop-

ulations, frequently only 2,3b in populations from the southeastern United States.

Ventral brush with 6 pairs of hairs; hairs 4b,4c-X usually 3,4b (2-4). Gills usually

tapering distally; dorsal usually much longer than ventral and relatively short, 0.9-

1.6 (0.5-2.0) length of saddle.

SYSTEMATICS. Aedes triseriatus can be readily separated from hendersoni and

brelandi by the following characters: in the adult by (1) the reduction in num-

ber and size or the absence of the anterior dorsocentral, humeral, lateral prescutal

and posterior fossal bristles, (2) the less extensive light scaling of the fossal area,

the light scales usually not extending as far mesad as the dorsocentral area, (3)

the claws of the foreleg and midleg, which are evenly curved and have a short

submedian tooth in the female, and which do not have an external tooth at the

base of the larger claw in the male, (4) the ornamentation of the wing, which

usually does not have white scales at the base of the costa in the female or on

the remigium (base of vein R) in the male, and (5) the fewer long bristles on

the palpus of the male; in the male genitalia most conspicuously by (1) the ab-

sence of a constriction in the tergal surface of the sidepiece, (2) the fewer elon-

gate setae of the basal tergomesal area of the sidepiece and their arrangement

in a narrow strip, (3) the setae on the mesal half of the middle portion of the

tergal surface of the sidepiece, which are not reduced in size or number, (4) the

smaller, more rounded sternomesal sclerite, (5) the moderately developed sterno-

mesal tuft with none or but very few of the setae strongly curved cephalad or

dorsad, and (6) the more evenly curved claspette filament; and in the larva by

(1) the presence of pigment in the thorax and abdomen, (2) the acus, which is

attached or detached but close to the base of the siphon, (3) the longer and usu-

ally more darkly pigmented pecten teeth, (4) the usually 4,5b (3-9) hair 1-X of

typical populations, (5) the ventral brush, which has 6 pairs of hairs and 4b,4c-X

usually 3,4b (2-4), and (6) the anal gills, which are unequal, with the dorsal longer

than the ventral, relatively short and tapered distally. The pupa of triseriatus can

usually be separated from that of hendersoni or brelandi by (1) the few branches

30 Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

in hair 1-IV, and (2) the shorter 5-IV.

There is a striking geographic variation in chaetotaxy and coloration of the

adults of triseriatus. Females from the southern two-thirds of Florida differ from

the typical triseriatus of the northeastern United States in the absence of most

anterior mesoscutal bristles and the great reduction of light scaling on the head,

mesoscutum, scutellum, haltere, femora and abdomen. Females from northern Flor-

ida, southern Georgia and coastal areas along the Atlantic northeast to North

Carolina and along the Gulf west to Louisiana usually also lack most of the an-

terior mesoscutal bristles and usually have the light scaling of the mesoscutum

and femora slightly reduced in extent. This partially darkened form seems to in-

tergrade with typical triseriatus to the north and even with the much darkened

form to the south. I have not been able to find any constant differences be-

tween the male genitalia, pupae or larvae of these darkened forms and northern

triseriatus. It is possible that the much darkened form from southern Florida is

a species distinct from triseriatus, but this cannot be determined at this time.

A great deal of additional collecting must be done in order to determine the

geographical and ecological distribution of these darkened forms and to docu-

ment the existence and extent of zones of intergradation between them.

The mouthparts of triseriatus larvae are quite variable. The maxillae may be

oval or triangular and the mental plate may be obtuse or acute. These variations

are correlated with each other and are frequently, but not always, correlated with

dimorphism of the mouthbrushes and antennae, as follows. Larvae with filamen-

tous mouthbrushes and long antennae usually have oval maxillae and an obtuse

mental plate, while those with pectinate mouthbrushes and short antennae fre-

quently have triangular maxillae and an acute mental plate. Larvae with the first

set of characteristics predominate in the northern portion of the range of ftriser-

iatus and larvae with the second set are more common in the southeastern United

States.

BIONOMICS. The immatures of triseriatus are usually found in treeholes and

are occasionally found in artificial containers and rockholes. They are frequently

associated with the immatures of hendersoni and zoosophus and rarely with those

of Ae. (Ochlerotatus) atropalpus. Adults have been collected at lights and in build-

ings and vehicles and females have been collected biting humans and horses.

Aedes triseriatus may be very important in the natural transmission of arbo-

viruses. It is a proven vector of the LaCrosse strain of California encephalitis virus

in Ohio and Wisconsin (Masterson et al., 1971:93-94,95; Sudia, Newhouse, Calisher

and Chamberlain, 1971:595-596; Thompson et al., 1972:94-95) and under labor-

atory conditions it is an effective vector of the viruses of yellow fever (Bennett

et al., 1939:101-105), eastern equine encephalomyelitis (Davis, 1940:49; Jakmauh,

1940:17; Chamberlain et al., 1954:280-283), Venezuelan equine encephalomye-

litis (Davis et al., 1966:227-230; Kissling and Chamberlain, 1967:68; Sudia, New-

house and Henderson, 1971:209), and western equine encephalomyelitis (Chamber-

lain et al., 1954:280-283).

DISTRIBUTION (fig. 1). Aedes triseriatus is found throughout the eastern United

States, from Minnesota to Maine, south to central Texas and Florida. The species

undoubtedly occurs in southeastern Canada and northeastern Mexico. Messersmith

(1971:7) reported finding 2 males of triseriatus in southern Greenland. The exis-

tence of a breeding population of this species there should be verified by finding

the immatures. Material examined: 4286 specimens; 1163 males, 1335 females,

1085 larvae, 703 pupae; 240 individual rearings (173 larval, 56 pupal, 11 incom-

plete).

Zavortink: New World Aedes 31

GREENLAND. Narssarssuag, 15 July 1957, D.H. Messersmith, 1 M gen [USNM].

UNITED STATES. Alabama: Auburn, Chewacla State Park, 27 Apr 1953, W.L. Seal, 2 L

[USNM]. Auburn, 8 Apr 1953, W.L. Seal, 1 L [USNM]. Ozark, Camp Rucker, 24 Feb-14 Mar

1943, J.G. Franclemont, 1 F [CDC], 39 M, 2 M gen, 24 F [CU]; 6 Mar 1944, 2 L [USNM];

J.G. Franclemont, 2 L [CU]. Sheffield, 7 June-18 Aug 1942, J.N. Belkin, 2 M, 1 M gen, 5 F

[UCLA]. Tuscaloosa, 19 Aug 1915, R.H. Van Ezdorf, 1 F [USNM]. Wilson Dam, Colbert Co.,

June-16 Aug 1954, R.X. Schick, 2 M, 2 M gen, 8 F, 10 P, 74 L [UCLA]. Wilson Dam, 27

July 1941-10 June 1942, J.N. Belkin, 1 pM (140), 1 pF (98), 1 M gen, 2 F [UCLA]; 14 Apr

1932-14 Apr 1937, 16 M, 1 F [CU]. Arkansas: Fayetteville, 8 May 1933, 1 M, 1 M gen [CU].

Helena, 30 July, H.S. Barber, 3 F [USNM]. Little Rock, Boyle Park, 28 Mar 1943, J.N. Belkin,

6 lp (441-446), 1 P [UCLA]. Little Rock, 14 Mar 1940, C.J. Carpenter, 1 M, 1 F [UTAH].

Scott, 1-10 June 1909, J.K. Thibault, 5 F [USNM]; 29 May 1914, D.L. VanDine, 1 F [USNM].

Stone Co., 5 Aug 1965, 1 F [USNM]. Connecticut: Fairfield Co., Aug 1910, V. Howard, 1 F

[USNM]. Redding, 14 Sept 1936, A.L. Melander, 1 F [USNM]. Delaware: Dover, 23-25 July

1935, 2 F [USNM]; 8-13 June 1944, 1 F, 14 L [CDC]. Newark, 8 Nov 1961-25 Mar 1963,

R.W. Lake, 1 M, 1 M gen, 1 F [USNM]. New Castle Co., 1962, 2 M, 1 F [UTAH]. Smyrna, 27

Aug 1933, D. MacCreary, 2 F [USNM]. Summit Bridge, 13 June 1963, R.W. Lake and F.W.

Kutz, 1 L [USNM]. District of Columbia: Catholic University, 12 Nov 1902-20 Sept 1906, T.

Pergande, 1 M, 9 F [USNM]. Chain Bridge, 26 June-Sept 1905, T. Pergande, 3 F [USNM].

Eastern Branch, 25 Aug, H.S. Barber, 1 F [USNM]. Site not specified, 4 Sept 1901, W.E. Hinds,

1 F [USNM]; May 1903, A. Busck, 1 F [USNM]; 15 June 1903, W.V. Warner, 1 M, 1 M gen,

1 F [USNM]; 2-11 Sept 1903, J. Kotinsky, 1 M, 2 F, 1 L [USNM]; 1 Aug 1904, T. Pergande,

5 M, 5 F [USNM]; 7 June 1908, H.O. Marsh, 1 F [USNM]. Florida: Avon Park, Aug 1943,

R.H. Arnett, Jr., 1 F [UCLA]. Gainesville, 11 Mar 1969, J.F. Reinert (69.20), 1 lpM; 17 Feb

1969 (69.21), 1 IpM; same data (69.22), 1 lpM; 21 Mar 1969 (69.23), 1 IpM; 29 Mar 1969

(69.29), 1 lpF; 19 Apr 1969 (69.35), 1 pM; 22 Apr 1969 (69.36), 1 pM; 19 June 1969 (69.40), 3

M, 4 F; 19-29 June 1969 (69.41), 6 M, 11 F; 28 June 1969 (69.44), 8 M, 2 F; 12 July 1969 (69.50),

8 M, 9 P; 30 July 1969 (69.57), 2 F, 2 P; 29 July 1969 (69.61), 3 M; 6 May 1969 (69.88), 8 M; 31

May 1969 (69.89), 12 M, 1 F; 11 Apr 1970 (70.42), 7 F, 7 P; 15 June 1970 (70.71), 11 IpF

(2-5,7-13), 3 pM (1,18,19), 5 pF (6,14-17); 17 June 1970 (70.72), 21 IpF (1-6,8-14,18,27,29-34),

4 pM (16,19,21,25), 8 pF (15,17,20,22-24,26,28), 1 F; 28 June 1970 (70.73), 2 IpM (27,41),

10 IpF (26,28-36), 15 pM (1-3,6,7,9,10,12,13,16-21), 7 pF (8,14,15,22-25), 4 M, 2 F, 6 P, 27 L;

29 Mar-28 July 1969, 43 M, 11 F [REINERT]. Indian River Co., 13 Nov 1970, G.F. O’Meara

(UCLA 578), 12 IpM (578-11,13,17-19,22,24,35 ,40,42,44,46), 26 IpF (578-10,12,15,16,20,21,

23,25-34,36-29,41,43,45,47,48), 7 pM (578-100-104,107,109), 3 pF (578-105,106,108), 1 Ip

(578-14), 13 M, 3 M gen, 6 F, 22 P, 30 L [UCLA]. Jacksonville, 14 Nov 1946, Miller, 1 F

[CDC]. Miami, 16,17 June 1934, J. Pearson, 4 F [CU]; Oct 1944, L. Stutz, 1 F [USNM];

8 Mar, 1 F [USNM]. Orlando, 30 July 1937, G.E. Quinby (FI-20), 3 M, 1 M gen, 1 F [USNM];

4 Aug 1937, G.E. Quinby (FI-27), 1 M, 1 M gen [USNM]; 6 Aug 1937, G.E. Quinby (FI-28), 1 L

[CDC]; 6 Apr 1968, L.T. Nielsen, 1 F [UTAH]; 2 Mar 1969, L.T. Nielsen, 2 M, 2 F [UTAH];

(1847), 1 M, 2 F; 16 June 1937 (1877), 2 M, 2 F; 23 June 1937 (1881), 2 L; 20 Jan 1936, 1 L

[USNM]. Ormond, 24 June 1940, 1 M gen [USNM]. Vero Beach, 18 Apr 1970, J.F. Reinert

(70.48), 5 L [REINERT]. West Palm Beach, 21 Mar 1947, Kaplan and W.F. Buren, 1 M, 1 M gen,

2 F [CDC]. No locality, 2 L [CDC]; (1499), 2 F; (1742), 2 M, 1 F; (1869), 1 F; (1885G1),

1 M; (2050), 1 F; (22-7), 1 M, 2 F; H.G. Dyar and A.N. Caudell (38), 3 M, 2 F [USNM].

Georgia: Atlanta, Fort McPherson, 2 Aug 1943, 3 F [UTAH]; 21 Mar-5 Apr 1944, 4M, 6 F

[USNM]; 6 July 1945, Gould, 1 L [CDC]; 23 June 1953, 1 L [CDC]. Atlanta, NW, 1-8 May

1947, 36 M, 1 M gen, 2 F [CDC]. Atlanta, 31 July 1906, W.B. Summerall, 2 F [USNM]; 29

July 1944, R.F. Fritz, 1 L [CDC]; 15 Aug 1944, H. Knutson, 2 L [CDC]; Apr 1946-31 May

1950, H.D. Pratt, 107 M, 94 F, 55 L [CDC], 2 M [UTAH]; 2 June 1947, R.C. Barnes, 2 F

[CDC]; 4 Mar-21 Apr 1948, C.J. Phillips, 5 F [CDC], 1 M [UTAH]; 23 Mar 1948, R.H. Foote

(436), 1 lpM (436-10), 8 IpF (436-2-4,7,9,11,12,16), 2 lp (436-6,8) [CDC]; 14 Feb 1949, H.D.

Pratt and C.J. Phillips, 13 L [CDC]; 3 May 1952, G. Heid, 1 F [UCLA]; 25 Mar 1953, W.D.

Sudia and R.H. Gogel, 13 M, 1 M gen, 1 F [CDC]; 1944, 2 M, 2 F, 1 L [USNM]; Mar 1948, 60 L

32 Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

[CDC]; 14 L [CDC]. Augusta Arsenal, Aug 1905, 1 M, 8 F [USNM]. Augusta, Camp Gordon,

26 Aug 1946, 4 L [CDC]. Billy’s Island, Okefenokee Swamp, June-7 July 1912, 1 M, 10 F [CU].

Black Rock Lake, 29 Apr 1946, 1 F [USNM]. College Park, 17 May 1948, R.H. Foote, 1 F

[CDC]. Columbus, Fort Benning, 10 Sept 1927, E.E. Hermes, 2 M, 1 F [USNM]; 3 May 1928,

E.E. Hume, 3 M, 1 F [USNM]; 15 Aug 1942, W.W. Middlekauff, 1 M gen [CU]; 1923,3M,7F

[USNM] ; 12 July 1924, 1 M [USNM]. Cornelia, 30 Mar 1948, 1 F [CDC]. Decatur Co., Spring

Creek, 14-29 July 1912, 2 F [CU]. Dublin, 13 Aug 1945, R. Dodge, 1 F [CDC]. Forsyth, 4 Mar -

1945, H. Knutson, 8 L [CDC]. Newton, Emory University, 8 Nov 1949, R.E. Bellamy, 1 M,

1 M gen [CDC]. Rabun Co., Black Rock Mountain, 20-25 May 1911, 2 F [CU]. Savannah,

10 Mar 1937, 1 M [USNM]. Union Co., Lake Trahlyta, 22 July 1945, Barrington, 2 L [CDC].

Worth Co., 31 Mar 1951, J.K. Wall, 2 L [CDC]. No locality, 22 Aug 1924, 2 F [USNM]; 5 M,

10 F [UTAH]. Illinois: Des Plaines, 30 June 1942, H.H. Ross, 1 M [UTAH]; 5 July 1942, H.H.

Ross and Mohr, 1 F [CDC]. Glencoe, July 1925, C.W. Edwards, 4 F [USNM]. Homer Park,

30 June 1927, 3 F [CU]. Indiana: Indianapolis, Fort Benjamin Harrison, 9 Aug 1923, 1 F

[USNM]. Lafayette, 13-24 July, J.M. Aldrich, 2 F [USNM]. Jowa: Ames, 17 Aug 1906, Quayle,

3 F [USNM]; 5 July-10 Sept 1919, 4 F [USNM]. Guttenburg (8.0 km N), 12 June 1967, M.

Manson, 4 L [UTAH]. Keosauqua, 5 Sept 1933, CCC Survey, 8 F [USNM]. Kansas: Franklin

Co., 30 Aug 1951, R.S. Roberts, 2 M, 1 F [UTAH]. Lawrence, May, H.T. Martin, 1 M, 1 F

[USNM]. Topeka, 21 June 1951, R.E. Beer, 2 M, 1 F [UTAH]. Kentucky: Brandenburg (6.4

km NE), 21 Aug 1941, G.E. Quinby and J.E. Rayl (Gq 465), 5 M, 3 F [USNM]. Corbin, 24

Aug 1904, H.S. Barber, 3 F [USNM]. Fort Knox (3.2 km S), 28 July 1941, G.E. Quinby

(Gq 439), 8 M [USNM]. Louisville, Cherokee Park, 9 Aug 1941, J.E. Rayl (Gq 484), 3 F

[USNM]; 22 Aug 1941, J.E. Rayl and G.E. Quinby (Gq 472), 3 F [USNM]. Louisville, 15

Aug 1935, 2 F [CU]. Louisiana: Alexandria, Camp Beauregard, 3 Apr 1943, W.W. Wirth, 1 M,

1 M gen [USNM]. Alexandria, 31 Mar 1943, W.W. Wirth, 1 F [USNM]. Baton Rouge, 15 Aug

1902-15 Aug 1903, 10 F [USNM]; 12 June 1941-5 Apr 1942, W.W. Wirth, 1 L [CDC], 1 F,4L

[USNM]. Buras, 1-8 June 1927, R.L. Turner, 1 F, 1 L [USNM]. Houma, 23-27 Apr 1931, 17M,

4M gen, 7 F [CU]. Jennings, 10 Oct 1906, A.A. Hammar, 1 F [CU]. Kilbourne, 13 May 1944-

18 Feb 1947, W.W. Wirth, 3 M [CDC], 1 F, 31 L [USNM]. Lafitte, 12 Apr 1931, 2 F [CU].

Lake Charles, USDA laboratory colony (UCLA 572), 10 IpM (572-20,22,25-30,32,34), 10 IpF

(572-21,23,24,31,33,35-39), 107 M, 57 F, 212 P, 175 L [UCLA]. Lake Charles, 14 July 1970,

H.C. Chapman (UCLA 573), 8 IpM (573-20-25,27,34), 9 IpF (573-26,28-33,35,36), 40 M, 3 M

gen, 17 F, 35 P, 10 L [UCLA]. Logansport, 23 Mar 1908, E.S. Tucker, 1 F [USNM]. Mound,

27 Apr 1915, D.L. VanDine, 4 M, 3 M gen, 2 F [USNM]; G.H. Bradley, 3 F [USNM]; 2 L

[USNM]. New Orleans, Camp Plauche, Apr-May 1944, 3 F [USNM]. Norco, 15 Oct 1944,

F.N. Young, 2 L [UCLA]. Port Jackson, 2M, 2F [USNM]. Rapides, 7 ‘Apr 1943, W.W. Wirth,

1 F [USNM]. St. Martinville, Longfellow Evangeline State Park, 8 Sept 1968, L.T. Nielsen

(N-33-68), 5 M, 3 F, 9 P, 7 L [UTAH]. Starks, 20 July 1970, H.C. Chapman (UCLA 575),

5 IpM (575-20,23,27,29,30), 6 IpF (575-21,22,24-26,28), 25 M, 1 F, 27 P, 36 L [UCLA]. No

locality, 3 M gen [USNM]. Maine: Naples, Sebago Lake State Park, J.C. Kuschke, 2 F [USNM].

Maryland: Baltimore, Patapsco State Park, 17 Oct 1965, W.A. McDonald (UCLA 286), 2 IpM

(286-30,31) [UCLA]. Baltimore, Nov 1922, W.A. Hoffman, 2 F [USNM]; 11 July 1919, 2 F

[UCLA]. Bethseda, 6 Aug 1949, K.L. Knight, 1 lpF (8.8), 1 M [UTAH]. Cabin John, 30 Aug-

5 Sept 1908, F. Knab, 2 M, 3 F [USNM]; 17 June 1928, H.G. Dyar, 1 M [USNM]. Camp Verde,

May-June 1919, R.C. Shannon, 1 F [USNM]. Edgewood, 11 Sept 1947, S.J. Carpenter, 1 F

[UTAH]. Forest Glen, 29 Aug 1966, W.W. Witth, 1 M, 1 M gen [USNM]. Herzog’s Island, 24

June 1906, 1 M [USNM]. Plummer’s Island, 5 June-19 Aug 1903, W.V. Warner, 2 F [USNM];

Sept 1903, A. Busck, 1 F [USNM]; 30 Aug 1908-28 July 1912, F. Knab, 2 F [USNM]; 13 Aug

1914, R.C. Shannon, 1 M [USNM]; 3 May 1936, H.S. Barber, 1 L [USNM]; 27 June 1965, W.

Grimm (UCLA 283), 1 IpM (283-10), 1 M gen [UCLA]; 25 Aug, R.P. Currie, 2 F [USNM];

R.P. Currie and F. Knab, 1 Ip (282) [USNM]. River View, Aug 1905, T. Pergande, 8 M, 1 M

gen, 3 F [USNM]. Rockville, 4 July 1960, C.W. Sabrosky, 1 F [USNM]. Silver Spring, 20

July 1940, F.C. Bishopp, 2 M [USNM]. Takoma Park, 1 F [USNM]. Massachusetts: Amherst,

1 June 1943, 2 M [CDC]. Athol, 10 Sept 1932, A.L. Melander, 1 F [USNM]. Ayer, Fort

Zavortink: New World Aedes 33

Devens, 25 July 1944, 1 M [CDC]. Chicopee, 17 Aug 1903, F. Knab, 1 F [USNM]. East Lee,

26 July 1921, J.L. Webb, 1 F [USNM]. Framingham, 20 Aug 1943, 6 F [CDC]. Granby, 12 Sept

1903, F. Knab, 1 F [USNM]. Springfield, W.H. Chaplin, 2 M, 1 M gen, 1 F [USNM]. Westfield,

30 July 1903, F. Knab, 1 F [USNM]. West Springfield, 26 July-24 Sept 1903, F. Knab, 3 M,

1 M gen, 20 F, 2 P [USNM]. Michigan: East Lansing, 21 June 1942, C.W. Sabrosky, 1 M, 1 F

[USNM]. Nottawa, June 1942, C.W. Sabrosky, 2 F [USNM]. Ogemaw Co., 1,2 Aug 1959, R.R.

Dreisbach, 1 F [USNM]. Minnesota: Detroit Lakes, 8,9 Aug 1940, W.A. Hoffman, 2 F [USNM].

Minneapolis, 3 July, K. Taylor, 1 F [USNM]. Mississippi: Agricultural College, 11 May-14 July

1905, G.W. Herrick, 1 M, 2 F [USNM]. Clarksdale, 1 Aug 1904, H.S. Barber, 1 F [USNM].

Columbus, Columbus Army Air Field, 13 May 1943, S.J. Carpenter, 1 F [UTAH]. Corinth,

14 Aug 1904, H.S. Barber, 1 F [USNM]. Gulfport, 6 Apr 1944, 4 L [USNM]. Hattiesburg,

Camp Shelby, 18 Aug 1943, 1 L [CDC]; 7 Feb-7 June 1944, 16 L [USNM]. Jackson, 8 Aug

1904, H.S. Barber, 1 F [USNM]. Scott, 24 Apr 1915, M.B. Mitzmain, 1 F [USNM]. Sibley,

A. Fleming, 5 M, 2 F, 1 L [USNM]. West Point, 11 Aug 1904, H.S. Barber, 1 M, 1 F [USNM].

Missouri: Boone’s Lick, 3 May 1966, C. Childers, 1 L [USNM]. Columbia, 20 Aug 1941, W.M.

Gordon, 2 F [CU]; 18 Aug, K.C. Sullivan, 1 F [USNM]. Eagle Rock, Roaring River State Park,

5 July 1942, A.B. Gurney, 1 F [USNM]. Jefferson Bks’., 21 Oct 1926-24 May 1927, E.C.

Bingham, 2 F [USNM]. Kansas City, July 1951, 1 M [CDC]. Neosho, Camp Crowder, 19 June-

27 July 1942, A.B. Gurney, 1 M, 1 M gen, 3 F [USNM]. Oakville, 24 Sept 1941, W.M. Gordon,

1 F [CU]. St. Louis (3.2 km W), Sept-Oct 1904, A. Busck, 2 M, 5 F [USNM]. St. Louis Co.,

4 June 1940-28 June 1941, W.M. Gordon, 5 F [CU]. Sikeston, 27 Aug 1921, M.F. Boyd, 3 M,

2 F [USNM]. Webster-Groves, 15 July 1919-28 May 1920, Satterthwait, 20 M, 9 F [USNM].

New Jersey: Chester, 24 Sept, J.M. Aldrich, 1 F [USNM]. Hewitt, 18 June 1904, 1 F [CU].

Lahaway, 31 May-6 June 1903, J.T. Brakeley, 4 M, 3 F [USNM]; 8 June-2 Aug 1903, H.G.

Dyar, 4 M, 5 F, 3 P, 2 L [USNM]. Menlo Park, 26 June 1947, S.J. Carpenter, 1 F [UTAH].

New Brunswick, 29 Aug, J.M. Aldrich, 1 F [USNM]. Paterson, 7 Sept, J.M. Aldrich, 1 F [USNM].

Ridgewood, 2 July 1911, M.D. Leonard, 1 F [CU]. Stelton, 23 June 1944, 6 L [CDC]. New

York: Albany, 10 Aug 1907, 3 F [CU]. Cayuta Lake, June-July 1943, R. Matheson, 97 M,

2 M gen, 97 F, 3 L [CU]. Cranberry Lake, 24-26 June 1963, W.W. Wirth, 1 M [USNM]. Fort

Niagara, 13 July-23 Aug 1944, 9 F [CDC]. Ithaca, Buttermilk, 10 July 1928, 1 F [CU]. Ithaca,

Six Mile Creek, 6 May 1922, R.C. Shannon and West, 4 L [CU]; 7 Sept 1922, R.C. Shannon,

1 F [CU]. Ithaca, Woodson Hill, 24 June 1914, 2 F [CU]. Ithaca, 1901-1903, O.A. Johannsen,

13 M, 8 F, 3 P, 4L [USNM]; Apr-May 1942, J.N. Belkin, 1 M, 1 M gen, 5 P [UCLA]; 4 July 1969,

G. Steyskal, 3 F [USNM]; 3 Aug 1901, 1 F [CU]; 20 July 1914-7 Sept 1942, 34 M, 132 F

[CU], 1 F [UCLA], 2 L [USNM]; 4 M, 12 F, 4 L [CU]. Manhattan, 2 Sept 1909, 1 F [CU].

McLean, 27 June 1940, 1 F [CU]. New York City, 15 Pine St., 27 July 1944, C. Slemrod, 1

M [CDC]. Niagara Falls, 24 July 1944, 1 F [CDC]. North River, 6-16 Aug 1950, C.W. Sabrosky,

1 F [USNM]. Orangeburg, Camp Shanks, 23 June 1944, 1 F [CDC]. Rome, 10 Aug 1944, 1 F

[CDC]; 19 May-3 Aug 1945, 4 F [CDC]. Suffolk Co., July-Aug 1928, 7 F [CU]. Syracuse, 28

June 1932, 1 F [CU]. Tannersville, 16 Sept 1926, L.O. Howard, 1 F [USNM]. Watkins Glen,

7 July 1928, 4 F [CU]. West Point, 2 Aug 1944, 1 F [CDC]. Yaphank, July 1909, W.T. Davis,

1 F [CU]. North Carolina: Asheville, 30 May 1944, H.F. Schoof, 5 L [CDC]. Bodie Island,

23 Sept 1960, 1 F [CDC]. Chapel Hill, 1 Apr 1970, R.L. Knight (UCLA 569), 6 IpM (569-

20-25), 1 lpF (569-26) [UCLA]; same data (UCLA 570), 1 IpF (570-90) [UCLA]. Charlotte,

1916, H.P. Barret, 1 F [USNM]. Flat Rock, 4 July 1914, A.H. Jennings, 1 M, 1 F [USNM];

22 July 1914, R.H. VanEzdorf and J.A. LePrince, 1 F [USNM]. Maxton, 24 June-13 Sept

1943, A.B. Klots, 1 M, 2 F [USNM]. Old Fort, 28 Sept 1933, CCC Survey, 3 F [USNM]. New

River, May 1942, G.E. Bohart, 1 L [UCLA]. Pinnacle, Aug 1906, F. Sherman, 1 F [USNM].

Roanoke Rapids, 20 Aug 1915, T.H.D. Griffiths, 1 F [USNM]. Wilmington, 1920, H.P. Barret,

1 M [USNM]. Ohio: Columbus, 18-28 May 1943, H.W. Smith, 2 M, 2 M gen, 1 F [UCLA].

Fairfield Co., 31 July 1951, F. Mead, 1 F [CDC]; 1 F [CDC]. Ravenna, 8 June 1970, M.J.

Zavortink (UCLA 571), 3 IpM (571-20,23,31), 4 IpF (571-22,27,39,41), 1 Ip (571-42), 2 M

gen [UCLA]. Toledo, 24-30 June 1915, C. Fox, 2 F [USNM]. Wilmot, 5 July 1965, T.J.

Zavortink (UCLA 444), 1 IpM (444-10), 6 IpF (444-11-16), 35 M, 2 M gen, 19 F, 94 P, 172 L

34 Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

[UCLA]. Oklahoma: Chickasha, 18 May 1944, 1 L [CDC]. Flint, 19 June 1937, Standish-

Kaiser, 1 M, 1 M gen [CU]. Stillwater, 21 Sept 1964-18 Apr 1965, J.F. Reinert, 4 L [REINERT].

Pennsylvania: Allentown, 22 July 1945, R.C. Barnes, 4 F [CDC]. Mt. Gretna, 4 M, 2 F [USNM].

Turtle Creek, 31 Aug 1947, 10 L [CDC]. South Carolina: Charleston, Naval Ammunition Depot,

6 July 1944, 1 L [CDC]. Columbia, 1 Aug 1906, 2 F [USNM]. Myrtle Beach, 3 July 1943

(L-24), 4 M [UTAH]; 10 July 1943-31 Mar 1944, 8 L [USNM]; 2 F [USNM]. Orangeburg Co.,

25 Feb 1944, 2 L [CDC]. Richland Co., 26 May 1944, 1 F [CDC]. Santee-Cooper Reservoir,

27 May-1 June 1945, C.W. Sabrosky, 27 M, 4 F [USNM]. Swansea, 6 Aug 1911, F. Knab, 1 F

[USNM]. Tennessee: Athens, 21,22 Aug, H.S. Barber, 1 M, 2 F [USNM]. Decherd, 18 Aug

1904, H.S. Barber, 1 F [USNM]. Kingsport, 18 July-18 Aug 1934, L. Arnott, 6 M, 1 M gen, 2 F,

1 L [CU]; 18 July 1934, W. Heinrich, 12 M, 2 M gen [CU]. Knoxville, 26-30 Apr 1935, S.E.

Shields, 1 M, 9 F [USNM]. Norris, 28 May 1935, S.E. Shields, 1 M [USNM]. Reelfoot Lake,

1 Sept 1941, M. Gordon, 2 F [CU]. Texas: Austin, 17 June 1947, O.P. Breland, 2 F [CDC];

30 July 1947-15 June 1948, O.P. Wilkins, 2 L [CDC]. Bexar Co., 20 July 1942, 1 M [CDC].

Brazoria Co., 2 Sept 1943, 1 L [CDC]. Brownsville, 15 May 1940, 1 F [CU]. Cyprus Bayou,

23 Aug 1903, J.D. Mitchell, 5 F [USNM]. Denison, 22,25 June 1904, H.S. Barber, 2 F [USNM].

Eagle Lake, 8 Sept 1968, L.T. Nielsen (N-34-68), 7 M, 17 F, 9 L [UTAH]. Frio Co., 2 June

1943, 1 M, 1 F [CDC]. Harris Co., 4 Aug 1943, 1 L [CDC]. Hondo, 9 Sept 1968, L.T. Nielsen

(N-35-68), 8 M, 15 F, 2 P, 9 L [UTAH]. Huntington, 24 Apr 1934, CCC Survey, 1 F [USNM].

Luling, Palmetto State Park, 11 Jan 1948, O.P. Breland, 2 L [CDC]. McAllen, 4 Nov 1925,

R.L. Turner, 1 F [USNM]. San Antonio, Fort Sam Houston, 13 Mar 1964, J.F. Reinert, 3 L

[REINERT]. San Antonio, 29,30 Apr 1942, W.C. Reeves, B. Brookman and R.B. Eads, 1 M, 1 F

[USNM]. Victoria, 28 July-18 Oct 1904, E.G. Hinds, 3 M, 7 F, 5 P, 4 L [USNM]. No locality,

1953, F.C. Harmston, 3 M, 2 F [UTAH]. Vermont: Jacksonville, Laurel Lake, 18 May 1947,

H.D. Pratt, 1 M, 7 F [CDC]. Virginia: Accotink, Fort Humphreys, 23 June 1921, 11 M, 1 M gen,

9 F [USNM]. Bluemont, 27-29 July 1904, 6 M, 3 F [USNM]. Difficult Run, 11 July 1906,

H.S. Barber and F. Knab, 1 F [USNM]. Falls Church, Holmes Run, 10 July 1960-17 Sept 1961,

W.W. Wirth, 10 M, 1 F [USNM]. Falls Church, 4 Sept 1906, A.N. Caudell, 1 F [USNM].

Glencarlyn, 10 May 1920, C.T. Green, 1 M, 1 F [USNM]. Great Falls, 3 Oct 1903-3 Sept 1909,

T. Pergande, 15 M, 24 F [USNM]; 15 June 1953, G. Steyskal, 1 F [USNM]; 22-28 June 1911,

3 M, 2 F [USNM]. Hampton, Fort Monroe, 19 July 1927, E. Slackshear, 1 F [USNM]. Lake

Drummond, 8-11 June 1905, H.S. Barber, 1 F [USNM]. Monterey, 20 Aug 1934, S.S. Cook,

1 M, 5 F [USNM]. Mt. Vernon, 15 July 1923, 1 F [USNM]. Richmond, July 1927, 1 F [CDC].

Rosslyn, 19 Sept 1902-July 1910, T. Pergande, 8 M, 11 F [USNM]; 20 Aug 1913, F. Knab, 1 F

[USNM] . Williamsburg, Camp Peary, July 1943, R.M. Bohart, 1 L [USNM]. Woodstock, 2 June

1903-12 Aug 1904, F.C. Pratt, 9 F, 8 P, 11 L [USNM]. No locality, 18 June 1903-5 Oct 1905,

T. Pergande, 4 M, 2 F [UCLA], 48 M, 78 F, 1 P [USNM], 2 F [UTAH]. Wisconsin: Darlington,

2 July 1950, R.E. Ryckman, 1 F [USNM]. State Unknown: New Smyrna, 23 July 1941, 1 M,

1 M gen [USNM].

LOCALITY UNKNOWN. 5 M, 2 F [CDC]; 1 IpF, 9 M, 8 M gen, 7 F, 2 L [CU]; 15 Aug

1919, 2 F; EJ. Gerberg (UCLA 284), 1 IpM (284-11), 1 IpF (284-10), 1 M gen; 1 M gen

(550310-22) [UCLA] ; May 1905, F. Knab (224), 2 M, 1 F; Marcovitch, 1 F; 1 M, 1 F [USNM].

ZOOSOPHUS GROUP

4. Aedes (Protomacleaya) zoosophus Dyar & Knab

Figs. 10-12

1918. Aedes zoosophus Dyar and Knab, 1918:165. TYPE: Holotype female, Kerrville, Texas,

United States of America, 19 Aug 1909, F.C. Pratt [USNM]. Synonymy with fluviatilis

(Lutz, 1904) by Dyar (1921a:30); resurrected by Dyar (1928:220-221).

Zavortink: New World Aedes 35

1925. Aedes alleni Turner, 1924:84. TYPE: Lectotype male with genitalia slide, Mission, Texas,

United States of America, reared from larva found in treehole, 28 Dec 1923 [USNM;

selection of Stone and Knight, 1956:214]. Synonymy with zoosophus by Gjullin (1946:

234).

Aedes (Finlaya) zoosophus of Gjullin (1946:234); Knight and Marks (1952:538,567); Carpenter

and LaCasse (1955:259-261); Stone, Knight and Starcke (1959:173); Carpenter (1968:83;

1970:54).

Aedes (Taeniorhynchus) zoosophus of Dyar (1928:220-221).

Aedes (?Taeniorhynchus) zoosophus of Dyar (1918:74).

Aedes (?Ochlerotatus) zoosophus of Edwards (1932:138).

Aedes (Finlaya) alleni of Dyar (1928:222-223); Edwards (1932:152); Matheson (1944: 185-186).

Aedes alleni of Dyar (1924:131-132); Rozeboom (1942:30); Jenkins and Carpenter (1946:36);

Breland (1949:93-100).

Aedes (Taeniorhynchus) fluviatilis in part of Dyar (1921a:30; 1922b:86-87; 1925:146).

FEMALE. Wing: 3.02 mm. Proboscis: 1.92 mm. Forefemur: 1.68 mm. Abdo-

men: about 2.8 mm. Head: Integument brown to black. Erect scales usually uni-

colorous, white to yellowish. Narrow curved scales usually unicolorous, silver-white.

Thorax: Integument brown to black. Acrostichal, anterior dorsocentral, humeral

and posterior fossal bristles usually absent, rarely a total of 1 or 2 present and

well developed; usually 1 well developed lateral prescutal bristle present. Meso-

scutum with background of predominantly small creamish-tan to golden-brown

scales anteriorly and dark brown or black scales posteriorly and with pattern of

predominantly larger silver-white or cream-colored scales, as follows: (1) cream-

colored scales in broad patch on median anterior promontory, (2) cream-colored

scales sometimes forming narrow broken or complete acrostichal line, (3) silver-

white scales in broad lateral prescutellar line, (4) silver-white scales usually in nar-

row to broad posterior outer dorsocentral line, and (5) cream-colored and silver

scales in broad to very broad lateral marginal line extending from anterior pro-

montory to wing root and mesad along scutal suture, the scales more distinctly

cream-colored anteriorly and more distinctly silver posteriorly. Midlobe of scu-

tellum with scales very dense, broad and flat, silver. Pon scales narrow and curved,

creamish-tan, yellowish-cream and/or cream-white. Pleural scale patches silver-white;

psp scales absent or few; ssp scales present; pst and metameron bare. Legs: Pos-

terior or ventral surface of forefemur and midfemur with light scales restricted

to basal 0.5-0.6. Knee spot absent to small on foreleg, small to moderate on mid-

leg, moderate to large on hindleg. Tibiae usually with basal light patch or ring

and sometimes with pale streak on posterior surface. Tarsal segments 1-3 or 4

of foreleg and midleg and tarsal segments 1-4 or 5 of hindleg with conspicuous

broad basal white ring. Claws of hindleg simple. Wing: Base of costa with white

scales in large patch or short line extending about 0.5 distance to crossvein h.

Base of vein R white scaled to slightly distad of level of crossvein h. Abdomen:

Tergites II-VIII usually dark scaled with basal light band that is broader laterally.

Basal sternites light scaled, distal dark scaled with broad basal light band that

may be broader laterally.

MALE. Essentially as in female except for sexual characters. Head: Palpus us-

ually with light scales on dorsal surface of segments 2 and 3; with numerous long

bristles from apex of segment 3 distad. Thorax: Mesoscutum with greater pro-

portion of scales of anterior portion silver-white to cream-colored than in female.

Psp scales usually absent. Legs: Larger claw of foreleg and midleg with an acute

basal external tooth; claws of hindleg simple. Wing: Remigium (base of vein R)

36 Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

light scaled to slightly beyond level of crossvein h.

MALE GENITALIA (fig. 11). Sidepiece: Usually weakly to moderately pig-

mented. Basal tergomesal area moderately to conspicuously swollen; setae of basal

tergomesal area few, not arising from a differentially sclerotized plaque; median

sternomesal sclerite weakly developed; median sternomesal tuft very weakly de-

veloped, the setae not strongly curved dorsad. Claspette: Filament evenly curved.

Phallosome: Aedeagus relatively long and slender, length 2.1-2.4 greatest width.

PUPA (fig. 11). Abdomen: about 3.3 mm. Trumpet: 0.48 mm. Paddle: 0.66

mm. Hairs largely strongly developed and strongly pigmented. Cephalothorax: Hairs

4.8-C moderately to strongly developed, short; 8-C shorter and finer than 9-C.

Trumpet: Medium golden-brown to brown. Abdomen: Hair 1-I] with 2-4 mod-

erately to strongly developed short branches; 1-III usually double (1-4b); hair 1-

IV usually double (1-3b); hair 1-V single or double (1-3b). Hair 2-II] mesad or

laterad of 1-IIl. Hair 4-Il mesad of 5-II. Hair 5-IV 1.8-2.6 length of 3-III. Hair

6-I,II strongly developed, as stout as or stouter than 3-[,II. Hair 9-III-VI stout,

relatively long. Paddle: Apex rounded or subtruncate.

LARVA (fig. 12). Head: 1.05 mm. Siphon: 0.85 mm. Anal Saddle: 0.32 mm.

Head: Labrum more or less evenly rounded in dorsal aspect. Hair 1-C stout, aris-

ing on or near front edge of labrum. Hair 4-C well developed, with numerous

(10-24) long branches; closer to midline than to 6-C. Hair 5-C single or double.

Hair 6-C usually 2,3b (2-4). Hair 7-C 6-9b. Hair 15-C short to moderately long,

1-3b. Mental plate distinctly triangular, lateral teeth not conspicuously enlarged.

Antenna: Shaft smooth or with a few spicules. Thorax: Pigment absent, living

larva white in color. Alveolar plate of hairs 1-3-P usually well developed. Hairs

1,5-P branched. Abdomen: Hair 12-I usually absent. Segment VUI: Comb scales

usually 10-12 (8-12), in 1 regular or irregular row. Siphon: Index usually 2.0-

2.3 (1.7-2.5). Acus attached or detached but near base of siphon. Pecten teeth

moderately long, usually darkly pigmented; usually 19-23 (11-24). Hair 1-S usu-

ally 2,3b (1-3). Anal Segment: Saddle large, extending 0.65-0.80 down lateral

surface of segment; with a narrow lightened or transparent ventral marginal or

submarginal area. Hair 1-X located along posterior margin of saddle about mid-

way between dorsal surface and ventral edge, its alveolus on saddle and removed

from edge of sclerotization by 2-4 alveolar diameters; usually 4,5b (2-7). Ventral

brush with 7 pairs of hairs; hairs 4b,4c-X usually 3b (2-4). Gills sausage-shaped;

dorsal much longer than ventral and moderately long, usually 1.5-2.5 (1.3-2.8)

length of saddle.

SYSTEMATICS. Aedes zoosophus is characterized as follows: in the adult by

(1) the creamish-tan to golden-brown background scales of the anterior half of

the mesoscutum, (2) the conspicuous white band at the base of tarsal segments

1-3 or 4 of the foreleg and midleg and 1-4 or 5 of the hindleg, and (3) the basally

banded tergites; in the male genitalia by the combination of (1) the weak to

moderate pigmentation, (2) the setae of the basal tergomesal area, which are few

and which do not arise from a differentially sclerotized plaque, (3) the weakly

developed sternomesal tuft, and (4) the relatively long, slender aedeagus, which

is 2.1-2.4 times longer than its greatest width; in the pupa by the combination

of (1) the short hairs 4,8-C, and (2) the relatively strong 6-I,J] and 9-III-VI; and

in the larva by (1) the absence of hair 12-I, and the combination of (2) the ab-

sence of pigment in the thorax and abdomen, (3) the few comb scales in | reg-

ular or irregular row, (4) the large anal saddle with a narrow ventral marginal

or submarginal lightened or transparent area, and (5) the ventral brush which has

Zavortink: New World Aedes Bi

7 pairs of hairs and 4b,4c-X usually 3b (2-4).

Aedes zoosophus differs so conspicuously from other species of Protomacleaya,

particularly in ornamentation of the adult and features of the larva, that I am

placing it into a monotypic group. Since it possesses a combination of primitive

characters, such as the undifferentiated genitalia of the male and the well developed

ventral brush and anal saddle of the larva, and derived characters, such as the

unusual ornamentation of the adult and the absence of hair 12-I in the larva,

zoosophus is probably an early segregate of Protomacleaya.

Too few specimens of research quality have been available for study to deter-

mine the existence or extent of geographical variation in zoosophus. The light

scaling of depauperate adults is reduced in extent and in depauperate males the

light scales of the palpus may be completely absent.

BIONOMICS. The immatures of zoosophus have been collected in treeholes and

artificial containers. They are frequently associated with 2 species of Protomacleaya

in the Triseriatus Group, hendersoni and triseriatus, and rarely associated with Ae.

(Ochlerotatus) epactius. Adults of both sexes are attracted to lights and females

are known to bite humans. |

DISTRIBUTION (fig. 10). Aedes zoosophus is definitely known from only the

southcentral United States and northeastern Mexico. Material examined: 166 spec-

imens; 40 males, 46 females, 64 larvae, 16 pupae; 12 individual larval rearings.

MEXICO. Tamaulipas: Ciudad Victoria, 30 Apr 1962, 1 L [USNM].

UNITED STATES. Arkansas: Fort Smith, Camp Chaffee, May 1964, Montague, 1 F [USNM].

Pine Bluff, 15 May 1964, Castillion, 1 F [USNM]. Kansas: Salina, T.A. Olson, 1 M gen [USNM].

Oklahoma: Cheyenne, 7 June 1937, Standish-Kaiser, 1 M, 1 M gen [CU]. Muse, 25 June 1937,

Standish-Kaiser, 1 F [CU]. Woodward (9.7 km E), Boiling Springs State Park, 1 Sept 1968,

L.T. Nielsen (N-32-68), 1 M, 1 F, 1 L [UTAH]. Texas: Austin, Zilkes Park, 23 Feb-10 Mar

1956, Steen, 2 M, 2 M gen, 6 L [UCLA]. Austin, 1941, C.P. Coogle, 1 F [USNM]. Brownsville,

29 Feb 1924, R.L. Turner, 1 F [USNM]; 25 Mar 1940, F.W. Fiske, Jr., 1 M, 1 M gen, 1 F

[USNM] ; 29,29 Sept 1942, E.S. Ross, 1 M, 2 F, 3 L [USNM]; 12 Sept 1944, C.R. Joyce, 23 L

[USNM]; 7 May 1963, J.F. Reinert, 2 L [REINERT]; 11-25 Mar 1940, 2 F [USNM]; 15 Apr-

15 May 1940, 4 F, 4 L [CU]. Corpus Christi, Naval Air Station, 3 June 1942, W.M. Gordon, 1 M,

1 M gen [CU]. Fort Worth, 31 May-8 June 1944, J.E. Porter, 2 M, 2 F [USNM]. Frio Co., 2

June 1943, 3 M, 3 F [USNM]. Hondo, 9 Sept 1968, L.T. Nielsen (N-35-68), 4 P, 4L [UTAH].

Mission, Dec 1923-Mar 1924, R.L. Turner, 10 M, 8 F, 5 L [USNM]. Rio Grande Valley, Oct

1925, R.L. Turner, 1 F [USNM]; 1 M [USNM]. San Antonio, Fort Sam Houston, 29 July

1964, J.F. Reinert, 2 L [REINERT]. San Antonio, 29 May 1941, 1 M [USNM]. San Benito,

6 June 1942, R.B. Eads, B. Brookman and W.C. Reeves, 3 M, 2 F [USNM]. Sheffield, 7 Mar

1949, A.G. Flury, 3 M, 3 M gen, 5 F [UCLA]; 30 May 1949, A.G. Flury, 3 M, 4 F [USNM]:

3 Sept 1969, T.J. Zavortink and J.A. Bergland (UCLA 619), 4 IpM (619-10,12,13,16), 5 IpF

(619-11,14,15,17,18), 1 M gen, 1 L; same data (UCLA 632), 1 lpF (632-10); same data (UCLA

633), 2 IpM (633-90,91), 1 M gen [UCLA].

Additional Record From the Literature

UNITED STATES. Louisiana: (see Carpenter, 1968:83).

KOMPI GROUP

FEMALES. Head: Integument brown to black. Erect scales usually bicolorous,

white to amber mesally, dark brown to black laterally. Narrow curved scales us-

ually bicolorous, white mesally, white and dark brown to black laterally. Thorax:

Integument brown to black. Acrostichal bristles numerous, moderately to strongly

38 Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

developed; anterior dorsocentral bristles numerous and well developed, reduced

in number and size, or absent from disc; humeral, lateral prescutal and posterior

fossal bristles few to numerous, weakly to strongly developed. Mesoscutum with

background of mostly small dark brown to black scales and with pattern of mostly

larger white to silver-white scales, as follows: (1) narrow to broad complete acros-

tichal line present or absent, (2) lateral prescutellar line usually present, (3) com-

plete posterior outer dorsocentral line usually present, (4) narrow to broad lat-

eral marginal line extending from anterior promontory to wing root developed,

and (5) posterior fossal line joining lateral marginal and posterior outer dorso-

central lines usually present. Midlobe of scutellum with scales sparse, narrow and

curved, white to silver-white or some dark. Ppn scales varied in shape, usually

white to silver-white; psp scales usually present, few to numerous; ssp scales pres-

ent; pst and metameron bare or scaled. Legs: Tarsal segment 1 of foreleg and

midleg marked with light scales, the marking varying from a relatively incon-

spicuous streak on basal portion of posterior and ventral or dorsal surfaces to

a conspicuous white band covering nearly entire segment; segment 2 of foreleg

and midleg and segment 1 of hindleg sometimes marked with white at base. Claws

of hindleg simple. Wing: Base of costa and vein R usually without light scales.

Abdomen: Tergites II-VIII dark scaled with basolateral light patch. Basal sternites

frequently light scaled, distal dark scaled with basolateral light patch or light basal

band which is broader laterally.

MALES. Essentially as in females except for sexual characters and usually

mesoscutal ornamentation. Head: Palpus entirely dark scaled; with few to num-

erous long bristles from apex of segment 3 distad. Thorax: Mesoscutum more ex-

tensively light scaled than in females, entirely white or silver-white in most species.

Legs: Larger claw of foreleg and midleg with an acute basal external tooth; claws

of hindleg simple. Wing: Remigium (base Of vein R) dark scaled or entirely or

partially light scaled to at least level of crossvein h. Abdomen: Tergites some-

times with conspicuous basal light band.

MALE GENITALIA. Sidepiece: Usually moderately to strongly lemented: Basal

tergomesal area slightly to moderately swollen; setae of basal tergomesal area few

to moderately numerous, not arising from a differentially sclerotized plaque; med-

ian sternomesal sclerite weakly to moderately developed; median sternomesal tuft

absent to moderately developed, the setae not strongly curved dorsad. Claspette:

Filament evenly curved. Phallosome: Aedeagus relatively short and broad, length

1.7-2.0 maximum width.

PUPAE. Hairs strong and deeply pigmented. Cephalothorax: Hairs 4,8-C very

strongly developed, long; 8-C much longer and stronger than 9-C. Trumpet: Light

golden brown to brown. Abdomen: Branches of hair 1-II moderately to strongly

developed, long. Hair 6-I,II usually strongly to very strongly developed, usually

as stout as 3-I,II. Hair 9-III-VI fine, short to relatively long. Paddle: Apex usually

pointed or produced.

LARVAE. Head: Labrum more or less evenly rounded in dorsal aspect. Hair

1-C stout, arising on or near front edge of labrum. Hair 4-C poorly developed,

with few (2-8) short branches; much closer to 6-C than to midline. Hair 5-C 1-4b.

Hair 6-C 1-7b. Hair 7-C 2-6b. Hair 15-C short to moderately long, 1-3b. Mental

plate distinctly triangular, lateral teeth not conspicuously enlarged. Antenna: Shaft

with numerous small spicules. Thorax: Pigment present. Hairs 1,5-P branched. Ab-

domen: Hair 12-I present. Segment VIII: Comb scales usually 24-55 (16-65), in

a patch. Siphon: Index 2.5-5.0. Acus usually attached, sometimes included in ba-

Zavortink: New World Aedes 39

sal sclerotization of siphon and not distinct. Pecten teeth 15-31. Hair 1-S 2-4b.

Anal Segment: Saddle moderate to large in size, extending 0.50-0.75 down lat-

eral surface of segment; without a narrow lightened or transparent ventral mar-

ginal or submarginal area. Hair 1-X 1-9b. Ventral brush with 6 pairs of hairs;

hairs 4b,4c-X usually 3,4b (2-5). Gills slender, tapered distally; dorsal subequal

to ventral or slightly longer, 1.1-3.5 length of saddle.

DISCUSSION. The Kompi Group is characterized as follows: in the adults by

(1) the bicolored head scales, and (2) the basal light marking on tarsal segment

1 of the foreleg and midleg; in the male genitalia by (1) the usually moderate

to strong pigmentation, and (2) the relatively short and broad aedeagus; in the

pupae by the strongly developed hairs 4,8-C; and in the larvae by (1) the patch

of usually numerous comb scales and the combination of (2) a weakly developed

hair 4C, and (3) 6 pairs of hairs in the ventral brush.

This group includes 6 species, burgeri, chionotum, kompi, niveoscutum, sandrae

and schicki. Each is differentiated from all the others by 1 or more novelties of

the adult and/or larva. The species are, in general, most easily and reliably sep-

arated as adults; the male genitalia are apparently indistinguishable. The 6 species

fall into 3 pairs, as follows: (1) kompi and burgeri, characterized by the usually

single hairs 5,6-C of the larva, the usually relatively few branched 1,2-I,II and

1,13-II-V of the larva and 1-III-V of the pupa, and the usual mesocaudal dis-

placement of 2-IV-VI of the pupa; (2) schicki and chionotum, characterized by

the presence of scattered light scales in the predominantly dark scaled areas of

the mesoscutum of the female, the conspicuous basal band on the abdominal

tergites of the male, and the usually multiple hairs 5,6-C on the head of the

larva; and (3) niveoscutum and sandrae, characterized by the predominantly dark

scaled midscutellar lobe in the female and the usually relatively highly branched

hairs 1,2-I,11 and 1,13-III-V of the larva and 1-III-V of the pupa in combination

with a relatively short siphon and paddle.

Most species of this group are characterized by strong sexual dimorphism in

adult ornamentation. In these, the mesoscutum of the male is completely or pre-

dominantly silver-white scaled while that of the female is extensively marked with.

dark scales.

The immature stages are found in treeholes, broken or cut bamboo internodes

and rockholes. Virtually nothing is known about the habits of the adults.

The species of the Kompi Group are found in mountainous regions from south-

eastern Arizona in the United States to western Guatemala. They are apparently

predominantly allopatric, separated either geographically or altitudinally.

KEYS TO SPECIES

FEMALES

ine Prosternum and metameron with well developed patch of scales; midlobe

of scutellum entirely or predominantly light scaled; dark scaled areas of

mesoscutum without scattered yellowish or whitish scales (figs. 13,14)

aR MA a lak ste ie Says ar eR yt ce MT Ue ke oT Mi 5. burgeri

Prosternum and/or metameron bare; if prosternum has patch of scales then

midlobe of scutellum predominantly dark scaled; if metameron has patch

of scales then dark scaled areas of mesoscutum with scattered yellowish

Or wWoitish scales toy te a eg oe le

40

2(1).

3(2).

4(3).

5(4).

2(1).

3(2).

Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

Metameron with patch of scales; base of hindtarsal segment 1 conspicuously

white scaled dorsally; dark scaled areas of mesoscutum with numerous

scattered yellowish or whitish scales (fig. 13) . . . . . . . .7. schicki

Metameron without definite patch of scales, sometimes 1 or more scales

present; base of hindtarsal segment | entirely dark scaled or with only

a few white scales dorsally; dark scaled areas of mesoscutum without

scattered yellowish or whitish scales except in chionotum ..... .3

Prosternum with extensive patch of scales; midlobe of scutellum predom-

inantly dark scaled; inner posterior fossal area without conspicuous white

patch <a, ASget oc tees . . 9. niveoscutum

Prosternum without definite patch of ‘scales: midlobe of scutellum predom-

inantly light scaled, or if predominantly dark scaled then inner posterior

fossal area with conspicuous wilitedcaled patel.) vec. te: im a's tae 0 od A

Erect scales of head very long and narrow, their width at apex scarcely

greater than width at midheight; dark scaled areas of mesoscutum with

numerous scattered yellowish or whitish scales (fig. 13) . . 8. chionotum

Erect scales of head moderately long and moderately broadened distally,

their width at apex 2-4 times greater than width at midheight; dark

scaled areas of mesoscutum without scattered yellowish or whitish scales

“3

Midlobe of scutellum predominantly light scaled; silver lateral prescutellar

scale line moderately broad, conspicuous; inner posterior fossal area with-

out conspicuous white scaled patch, dark scaled, or if white scaled then

these white scales broadly joined to lateral prescutal white line (fig. 13)

; ' . 6. kompi

Midlobe of scutellum predominantly dark ‘scaled: ‘silver lateral prescutellar

scale line usually narrow, inconspicuous; inner posterior fossal area with

conspicuous white scaled patch (figs 13) serdar 1Ossamadrae

MALES

Mesoscutum with numerous brown scales in conspicuous broad complete

dorsocentral line. . . . . . .6.kompi

Mesoscutum entirely silver-white scaled | or with relatively few brown scales

Tiahdttow (COUN ele GOrsOcedtital UNC. ae ket wanes ee ww

Prosternum and metameron with well developed patch of scales; palpus as

long as or longer than proboscis (fig. 14). . . ... . . . 5. burgeri

Prosternum and/or metameron bare; if either prosternum or metameron has

patch of scales then palpus shorter than proboscis . .........3

Metameron with patch of scales; base of hindtarsal segment 1 conspicuously

white scaled dorsally; palpus shorter than proboscis . . . . .7. schicki

Metameron without definite patch of scales, sometimes with 1 or more

scales; base of hindtarsal segment 1 entirely dark scaled, or if white scaled

dorsally then palpus subequal in length or or longer than proboscis. . .4

4(3).

5(4).

2(1).

3(2).

2(1).

3(2).

4(2).

Zavortink: New World Aedes 41

Prosternum with extensive patch of scales; remigium (base of vein R) dark

scaled; palpus usually shorter than proboscis . . . . . 9.niveoscutum

Prosternum without definite patch of scales; remigium predominantly light

scaled; palpus subequal in length to or longer than proboscis . . . . .5

Abdominal tergites with conspicuous white basal band . . . 8. chionotum

Abdominal tergites not banded or with inconspicuous light basal band

. 10. sandrae

PUPAE

Hair 1-II usually 6-8b (5-11) and paddle index usually 1.4-1.6 (1.3-1.7); hair

5-IV usually double (1-4b) (figs. 19,21) . . . . 7. schicki; 8. chionotum

Hair 1-II with fewer branches and/or paddle index less; 5-IV usually single

(single or double)

Hair 2-V usually slightly to greatly cephalad of level of 3-V; paddle index

usually 1.1-1.3 (1.1-1.4) (figs. 23,25) . . . 9. niveoscutum; 10. sandrae

Hair 2-V PERN caudad of level of 3-V; Pee index hie 1.4-1.5 Ma 2-

te: gigas nee f bia ; , , ; .3

Hair 10-C moderately developed, distinctly finer and shorter than 12-C;

most primary branches of float hair (1-I) usually with numerous second-

ary branches; branches of 1-II usually moderately developed and finer

than primary branches mE ES LG eae bet . . 5. burgeri

Hair 10-C strongly developed, usually distinctly stouter and longer than 12-

C; most primary branches of float hair (1-I) without secondary branches;

branches of 1-II strongly developed, as or nearly as stout as primary

branches of dl 4fie, 19)-.5 isc-0 thcen\t. anet Gos Bie m teeeceee komm

LARVAE

Apex of larger comb scales produced into moderate-sized spine; most caudal

hair of ventral brush (4a-X) short, ba mer oe 1.0-1.5 length of anal

Saqdle thin, 26) 0 °F Soy ; . . . 10. sandrae

Apex of larger comb scales evenly fringed | or a or 2 apical elements of

fringe slightly Cat hair 4a-X she passes at least 2.0 length of

anal saddle . . . ie may 9 5 ESBS iol OT i

Hair 6-C usually 3-5b (2-6); siphon index greater than3.4 ...... .3

Hair 6-C single or double and/or siphon index less than3.2. . .... .4

Branches of hairs 1,13-II-V relatively stout and coarse to near apex; 1-III-V

usually 3,4b (2-5); hair 5-V-VII usually single or double (1-3b), the

branches stout and coarse to near apex (fig. 20) . . . . . . .7.schicki

Branches of hairs 1,13-II-V finer and tapering from near base; 1-III-V usu-

ally 6,7b (5-10); i aie 5-V-VII usually 4,5b eur ae the branches finer and

tapering from near base (fig. 22). . . . . 8. chionotum

Hair 5-C usually double (1-3b); hair 2-II usually 2,3b (2-4); hair 1-II-V

eae a 3,4b hs ne rk index ftv nd 2.7-3.0 (2. 5-3.1) (fig. 24).

: 9. niveoscutum

42 Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

Hair 5-C usually single (single or double); 2-II usually single (1-4b); hair

1-II-V usually 2,3b (1-4); siphon index frequently greater than 3.1 (2.5-

SO iD

5(4). Branches of hair 1-VIII finer than those of 1-X (fig. 16). . . . 5. burgeri

Branches of hair 1-VIII stouter than those of 1-X (fig. 18) . . . .6.kompi

5. Aedes (Protomacleaya) burgeri Zavortink, n.sp.

Figs. 10,13-16

TYPES: Holotype male with associated larval and pupal skins (UCLA 562-47), Bodie Canyon,

13 air km east of Lochiel, Santa Cruz Co., Arizona, United States of America, elevation about

1530 m, egg from rothole in living sycamore tree, 27 Dec 1969, L.T. Nielsen and T.J. Zavortink

[USNM]. Allotype female with associated larval and pupal skins (UCLA 562-71), same data

as holotype [USNM]. Paratypes: 7 |pM (562-46,52,73,74,79,81,82), 20 lpF (562-44,50,53,54,

57-60,62-65 ,67,68,70,72,75-77,80), 1 pM (562-101), 1 pF (562-100), 12 M, 4 F, 1 M gen, 24 P,

272 L, same data as holotype (UCLA 562); 2 IpF (559-31,32), 1 M, 1 F,3 P, 64 L, same data as

holotype except eggs from rothole in living evergreen oak tree (UCLA 559); 4 lpM (567-20,

29,37,39), 7 IpF (567-23 ,24,26,32,34,36,40), 1 lp (567-38), 2 M, 1 M gen, 1 F, 9 P, 19 L,

same data as holotype except eggs from rothole in living evergreen oak tree (UCLA 567) [UCLA,

ISET, BM]; 20 M, 3 M gen, 20 F, 11 P, 14 L, same data as holotype except collected on 21 Mar

1968 by L.T. Nielsen, J.H. Arnell and J.H. Linam (HA-12-68) [UTAH]. This species is dedi-

cated to the original collector, John F. Burger.

Aedes n.sp. near kompi of Zavortink (1970:8); Arnell and Nielsen (1972:16).

Aedes (Finlaya) kompi of Burger (1965:396-398); Stone (1967:208).

FEMALE (figs. 13,14). Wing: 4.00 mm. Proboscis: 2.45 mm. Forefemur: 2.49

mm. Abdomen: about 3.3 mm. Head: Erect scales moderately long and moder-

ately broad, their width at apex 2-4 times greater than width at midheight. Tho-

rax: Anterior dorsocentral bristles numerous and well developed; humeral, lateral

prescutal and posterior fossal bristles few. Dark scaled areas of mesoscutum with-

out scattered yellowish or whitish scales. Light mesoscutal lines conspicuous; acros-

tichal line usually broad; lateral prescutellar line broad; posterior outer dorsocentral

line usually moderately broad; lateral marginal line very broad, usually extending

mesad to inner margin of fossa; posterior fossal line not broadened mesally and

forming a conspicuous more or less isolated patch. Midlobe of scutellum with

scales entirely or predominantly white. Most pleural patches dense and well de-

fined, consisting of broad flat to slightly outstanding imbricate silver-white scales.

Upper anterior ppn with scales moderately dense, most moderately broad to broad,

flat; pst with well developed oblique patch of scales; metameron with patch of

scales. Legs: Posterior surface of forefemur and midfemur with white streak usu-

ally confined to basal 0.5-0.6. Foretibia and midtibia entirely dark scaled or streak-

ed or speckled with white on posterior surface. Foretarsal segment | usually marked

with white on at least ventral and posterior surfaces in at least basal 0.6; mid-

tarsal segment 1 usually marked with white on all surfaces in at least basal 0.6;

hindtarsal segment 1 usually entirely dark scaled.

MALE (fig. 14). Essentially as in female except for sexual characters and meso-

scutal ornamentation. Head: Palpus usually longer than proboscis; with numerous

long bristles from apex of segment 3 distad. Thorax: Mesoscutum usually entirely

Zavortink: New World Aedes 43

silver-white scaled, rarely with a few dark scales in posterior inner dorsocentral

area. Wing: Remigium (base of vein R) entirely dark scaled. Abdomen: Tergites

not banded or with narrow light scaled basal band.

MALE GENITALIA (fig. 15). Apparently indistinguishable from other species

of group. |

PUPA (fig. 15). Abdomen: about 3.8 mm. Trumpet: 0.54 mm. Paddle: 0.82

mm. Cephalothorax: Hair 10-C moderately developed, distinctly finer and shorter

than 12-C. Abdomen: Float hair (1-1) moderately developed, with longest branches

not extending to lateral margin of segment I, and with most primary branches

usually with numerous secondary branches; branches of 1-II usually 3-5 (2-6), us-

ually moderately developed and finer than primary branches of 1-I; hair 1-III-V

usually 1-3b (1-4). Hair 2-IV-VI usually mesad of hair 1 of corresponding seg-

ment and usually relatively close to caudal margin of segment; 2-V usually caudad

of level of 3-V. Hair 5-IV single. Paddle: Index usually 1.4-1.5 (1.2-1.7).

LARVA (fig. 16). Head: 0.96 mm. Siphon: 0.99 mm. Anal Saddle: 0.38 mm.

Head: Hair 5-C usually single (single or double). Hair 6-C single. Hair 14-C usu-

ally single (single or double). Abdomen: Hair 1-II-V usually 2,3b (1-4), the branches

normal. Hair 2-II usually single (1-4b). Hair 5-II-VII usually 3,4b (2-4), the branches

normal. Hair 13-II usually 4,5b (3-6) and 13-III-V usually 2-4b (1-4), the branches

normal; 13-VI usually 6-8b (4-9), the branches much shorter and finer than those

of 13-VII; hair 13-VII 2-4b. Segment VIII: Branches of hair 1 finer than branches

of 1-X. Comb scales usually 24-38 (22-50); apex of larger scales fringed. Siphon:

Index about 3.1-3.9. Anal Segment: Hair 4a-X long, at least 2.0 length of anal

saddle.

SYSTEMATICS. Aedes burgeri is closely related to kompi and is distinguished

from that species as follows: in the adult by (1) the entirely silver-white scaled

mesoscutum of the male, (2) the more conspicuous light mesoscutal markings of

the female, and (3) the presence of scales on the prosternum and metameron in

both sexes; in the pupa by (1) the usual presence of numerous secondary branches

on the primary branches of the float hair (1-1), and (2) the more weakly devel-

oped 10-C and 1-II; and in the larva by the weaker branches of hair 1-VIII. This

species is the only one in the Kompi Group that has a well developed patch of

scales on both the prosternum and metameron of the adult.

Except for variation in development of the lateral prescutal, acrostichal and

posterior outer dorsocentral light scale lines in the female, burgeri is relatively

uniform in all stages throughout most of its range. However, both the larva and

pupa are extremely variable at the southernmost locality at which the species

has been found, where it occurs with niveoscutum. Here the larva may have more

highly branched hairs and a shorter siphon than is normal and the pupa may have

more highly branched or abnormally developed hairs and a shorter paddle than

usual. This variation is indicative of hybridization and introgression between burg-

eri and niveoscutum.

BIONOMICS. The immatures of burgeri are found in treeholes. In the north-

ern portion of its range this species has been collected in oak-pine forest, ripar-

ian woodland and scattered groves of oaks where it has been associated | or

more times with Ae. (Abraedes) papago, Ae. (Kompia) purpureipes, Ae. (Och-

lerotatus) monticola Belkin & McDonald, 1957 and Ae. (O.) muelleri. In the south-

ern portion of its range it has been collected in oak-pine forest where it has oc-

curred alone or in association with another species of the Kompi Group, niveo-

scutum. Burger (1965:396) reported finding adults of burgeri (as kompi) resting

on the sides of a treehole and on nearby leaves.

44 Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

DISTRIBUTION (fig. 10). Aedes burgeri extends from southeastern Arizona

in the United States to Jalisco in Mexico. In Arizona it has been collected at

elevations between 1070 and 1530 meters and in Mexico between 1400 and 1770

meters. Material examined: 1204 specimens; 145 males, 173 females, 614 larvae,

272 pupae; 109 individual rearings (106 larval, 2 pupal, 1 incomplete).

MEXICO. Jalisco: El Mirador (2.4 km S Quililla), 8 June 1971, T.J. Zavortink and L.T.

Nielsen (MEX 669), 1 lpM (669-10), 2 lpF (669-13,15), 1 M, 1 M gen, 2 P, 2 L; same data

(MEX 670), 1 M, 2 P, 3 L; same data (MEX 671), 1 lpM (671-13), 4 IpF (671-12,14-16), 4 M,

1 F, 5 P, 1 L; same data (MEX 675), 4 lpM (675-11-14), 6 IpF (675-15-17,19,22,23), 38 M,

1 M gen, 12 F, 56 P, 49 L [UCLA]; same data (N-22-71), 3 IpM (3,5,6), 6 IpF (7,9,10-13);

same data (N-23-71), 1 lpM (2), 1 lpF (1); same data (N-26-71), 2 IpM (1,2); same data (N-27-

71), 5 lpF (1-3,4,7) [UTAH]. Guadalajara (24 km NW), 7 June 1971, T.J. Zavortink and L.T.

Nielsen (MEX 667), 1 IpM (667-10), 4 lpF (667-11-14), 3 M, 1 M gen, 5 P, 13 L [UCLA].

Sinaloa: Potrerillos (5.7 km E), 11 June 1971, L.T. Nielsen and T.J. Zavortink (MEX 700), 8 lpM

(700-10-17), 4 lpF (700-18-21), 8 M, 2 M gen, 2 F, 15 P, 11 L [UCLA]; same data (N-51-71),

5 IpM (1-4,6), 1 lpF (5) [UTAH]. Sonora: Nogales (14.4 km S), 22 Aug 1970, J.H. Arnell and

L.T. Nielsen (HA-20-70), 7 M, 27 P, 50 L [UTAH].

UNITED STATES. Arizona: Bodie Canyon, Lochiel (13 km E), type series, see above. Men-

doza Canyon, Coyote Mountains, Pima Co., 23 Dec 1969, L.T. Nielsen (UCLA 550), 2 IpM

(500-30,34), 2 IlpF (550-31,32), 1 M gen, 9 L [UCLA]. Patagonia (3-6 km WSW), 18 Aug-

17 Sept 1964, J.F. Burger, 2 M, 2 M gen, 59 F, 2 P [UCLA], 3 M, 2 M gen, 7 F, 2 P [USNM]; 13

Sept 1968, T.J. Zavortink (UCLA 454), 1 IpM (454-12), 1 M, 1 M gen [UCLA].

6. Aedes (Protomacleaya) kompi Vargas & Downs

Figs. 10,13,17,18

1950. Aedes (Gualteria) kKompi Vargas and Downs, 1950:167-170. TYPE: Holotype female,

Tepoztlan, Morelos, Mexico, larva from treehole or rockhole, 29 June 1947, W.G. Downs

[ISET].

Aedes (Finlaya) kompi of Stone, Knight and Starcke (1959:165); Schick (1970a: 16).

FEMALE (fig. 13). Wing: 3.84 mm. Proboscis: 2.38 mm. Forefemur: 2.36 mm.

Abdomen: about 2.6 mm. Head: Erect scales moderately long, moderately broad,

their width at apex 2-4 times greater than width at midheight. Thorax: Anterior

dorsocentral bristles usually few and small or absent; humeral, lateral prescutal

and posterior fossal bristles few or absent. Dark scaled areas of mesoscutum with-

out scattered yellowish or whitish scales. Light mesoscutal lines relatively incon-

spicuous to conspicuous; acrostichal line usually present, narrow to moderately

broad; lateral prescutellar line moderately broad; posterior outer dorsocentral line

usually very narrow and incomplete; lateral marginal line narrow to very broad

and then extending mesad to inner margin of fossa; posterior fossal line very nar-

row to broad, not broadened mesally and forming a conspicuous more or less

isolated patch. Midlobe of scutellum with scales usually entirely or predominantly

light. Most pleural patches very dense and well defined, consisting of broad flat

to appressed imbricate silver-white scales. Upper anterior ppn with scales sparse

to moderately dense, moderately broad to broad, flat to appressed; pst usually

bare, rarely with scales near ppl; metameron bare. Legs: Posterior surface of fore-

femur and midfemur with white streak of basal portion extending to or near apex

as a narrow line. Foretibia and midtibia dark scaled or streaked or speckled with

white on posterior surface. Foretarsal segment 1 usually marked with white on

Zavortink: New World Aedes 45

at least ventral and posterior surfaces in at least basal 0.4; midtarsal segment 1

usually marked with white on all surfaces in at least basal 0.5; hindtarsal seg-

ment 1 dark scaled.

MALE. Essentially as in female except for sexual characters. Head: Palpus sub-

equal to proboscis in length or slightly longer; with relatively few long bristles

from apex of segment 3 distad. Thorax: Conspicuous broad complete brown scaled

dorsocentral line developed. Wing: Remigium (base of vein R) entirely dark scaled.

Abdomen: Tergites without basal band.

MALE GENITALIA (fig. 17). Apparently indistinguishable from other species

in group.

PUPA (fig. 17). Abdomen: about 3.2 mm. Trumpet: 0.55 mm. Paddle: 0.79

mm. Cephalothorax: Hair 10-C strongly developed, at least as stout and long as

12-C. Abdomen: Float hair (1-I) moderately to strongly developed, with longest

branches usually extending to lateral margin of segment I, and with most pri-

mary branches unbranched; branches of 1-II usually 2-5 (2-7), strongly developed

and as or nearly as stout as primary branches of 1-I; hair 1-IIJ-V usually 1-3b

(1-4). Hair 2-IV-VI usually mesad of hair 1 of corresponding segment and usually

relatively close to caudal margin of segment; 2-V usually caudad of or at the level

of 3-V, sometimes slightly cephalad of it. Hair 5-IV usually single (single or dou-

ble). Paddle: Index usually 1.4-1.5 (1.2-1.6).

LARVA (fig. 18). Head: 0.92 mm. Siphon: 0.79 mm. Anal Saddle: 0.33 mm.

Head: Hair 5-C usually single (single or double). Hair 6-C single. Hair 14-C us-

ually single (single or double). Abdomen: Hair 1-II-V usually 2,3b (1-3), the

branches normal. Hair 2-II usually single (1-3b). Hair 5-II-VII 2,3b, the branches

normal. Hair 13-II usually 3,4b (2-4) and 13-III-V usually 2,3b, the branches

normal; 13-VI usually 6-9b (5-11), the branches much shorter and finer than

those of 13-VII; hair 13-VII 2,3b. Segment VIII: Branches of hair 1 stronger than

branches of 1-X. Comb scales usually 35-55 (16-65); apex of larger scales fringed.

Siphon: Index usually 2.8-3.5 (2.5-3.9). Anal Segment: Hair 4a-X long, at least

2.0 length of anal saddle.

SYSTEMATICS. Aedes kompi is apparently most closely related to burgeri and

is distinguished from it as follows: in the adult by (1) the broad longitudinal

lines of dark scales on the mesoscutum of the male, (2) the more restricted light

mesoscutal markings of the female, and (3) the usual absence of scales on either

the prosternum or metameron; in the pupa by (1) the usual absence of second-

ary branches on most primary branches of the float hair (1-I), and (2) the more

strongly developed 10-C and 1-II; and in the larva by the stronger branches of

hair 1-VHI. Aedes kompi is conspicuously differentiated from all other species

of the group by the ornamentation of the mesoscutum of the male.

BIONOMICS. The immatures of kompi have been collected in treeholes and

holes in volcanic rock. In the former habitat they have been associated with 2

species of Aedes (Protomacleaya) in the Terrens Group, gabriel Schick, 1970 and

idanus Schick, 1970; in the latter habitat they have been found only with gabriel.

All known adults of kompi have been reared and nothing is known about their

habits.

DISTRIBUTION (fig. 10). Aedes kompi is presently known from a single area

along the Southern Escarpment of the Mesa Central in the State of Morelos, Mex-

ico, at an elevation of about 1600 meters. Material examined: 170 specimens; 18

males, 12 females, 100 larvae, 40 pupae; 28 individual rearings (13 larval, 10 pu-

pal, 5 incomplete).

46 Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

MEXICO. Morelos: Vicinity of Tepoztlan (including Gabriel Mariaca), 29 June 1947, W.G.

Downs, 1 M, 1 M gen, 1 F [USNM]; 7 Sept 1965, D.A. Schroeder (MEX 346), 3 pF (346-10,

100,101), 5 L; same data (MEX 349), 1 pM (349-100), 1 pF (349-101), 1 M gen, 4 L; same

data (MEX 350), 2 lpM (350-17,24), 1 IpF (350-25), 3 pM (350-93,106,112), 4 Ip (350-12,

30-32), 4 M, 3 M gen, 11 P, 72 L; same data (MEX 351), 2 pM (351-100,101), 1 M gen,

1 P, 1 L; same data (MEX 352), 2 IpM (352-12,17); 11 Aug 1970, K. and D.A. Schroeder

(MEX 619), 1 lpM (619-11), 5 lpF (619-16-19,25), 1 lp (619-27), 1 M; same data (MEX 620),

1 IpF (620-17); same data (MEX 624), 1 lpM (624-30) [UCLA].

7. Aedes (Protomacleaya) schicki Zavortink, n.sp.

Figs. 10,13,19,20

TYPES: Holotype male with associated larval and pupal skins (MEX 698-22), 4.8 km E of

La Emerta, Durango, Mexico, elevation about 2620 m, larva from rothole in living oak tree,

11 June 1971, L.T. Nielsen and T.J. Zavortink [USNM]. Allotype female with associated larval

and pupal skins (MEX 698-14), same data as holotype [USNM]. Paratypes: 11 lpM (698-10-

13,15-19,21,23), 1 lpF (698-20), 4 F, 3 M gen, 3 L, same data as holotype (MEX 698) [UCLA,

ISET, BM]; 1 lpM (4), 2 IpF (1,2), 3 pM (3,7,8), 1 pF (5), same data as holotype (N-50-71)

[UTAH]. This species is dedicated to Robert X. Schick in recognition of his contributions to

the knowledge of Protomacleaya.

FEMALE (fig. 13). Wing: 4.53 mm. Proboscis: 2.80 mm. Forefemur: 2.59 mm.

Abdomen: about 3.3 mm. Head: Erect scales moderately long and relatively nar-

row, width at apex 1.5-2.5 times width at midheight. Thorax: Anterior dorso-

central bristles numerous and well developed; humeral, lateral prescutal and pos-

terior fossal bristles numerous. Dark scaled areas of mesoscutum with numerous

yellowish or whitish scales. Light mesoscutal lines relatively inconspicuous; acros-

tichal line absent, narrow and broken, or moderately broad; lateral prescutellar

line narrow and broken to moderately broad; posterior outer dorsocentral line

narrow and broken to moderately broad; lateral marginal line narrow; posterior

fossal line narrow, sometimes broadened and denser mesally and then appearing

as a patch. Midlobe of scutellum entirely or predominantly light scaled. Most

pleural patches dense and moderately well defined, consisting of broad flat to

outstanding somewhat disheveled white scales. Upper anterior ppn with scales mod-

erately dense, narrow and curved to moderately broad and flat; pst sometimes

with scales near ppl; metameron with patch of scales. Legs: Posterior surface of

forefemur and midfemur with white streak usually extending from base to or

near apex. Foretibia usually conspicuously streaked with white from near base

to near apex on posterior surface; midtibia entirely dark scaled or posterior sur-

face speckled or streaked or white. Foretarsal segment 1 marked with white on

ventral and posterior surfaces in at least basal 0.6; midtarsal segment 1 marked

with white on all surfaces in at least basal 0.6; hindtarsal segment | with a con-

spicuous white streak or patch at base of dorsal surface.

MALE. Similar to female except for sexual characters and mesoscutal ornamen-

tation. Head: Palpus distinctly shorter than proboscis; with numerous long bristles

from apex of segment 3 distad. Thorax: Mesoscutum usually entirely silver-white

scaled, rarely with a few dark scales in posterior inner dorsocentral area. Wing:

Remigium (base of vein R) partially to entirely light scaled. Abdomen: Tergites

usually with conspicuous white scaled basal band.

MALE GENITALIA (fig. 19). Apparently indistinguishable from other species

in group.

Zavortink: New World Aedes 47

PUPA (fig. 19). Abdomen: about 4.2 mm. Trumpet: 0.72 mm. Paddle: 1.12

mm. Apparently indistinguishable from chionotum. Cephalothorax: Hair 10-C mod-

erately to strongly developed, usually longer and stronger than 12-C. Abdomen:

Float hair (1-I) moderately to very strongly developed, with longest branches some-

times extending beyond lateral margin of segment I, and with primary branches

with or without secondary branches; branches of 1-II usually 6-8 (5-11), strongly

developed and finer than or subequal in stoutness to primary branches of 1-I;

hair 1-III-V usually 3-6b (2-6). Hair 2-IV-VI usually in line with or laterad of

hair 1 of corresponding segment and usually relatively far from caudal margin

of segment; 2-V far cephalad of level of 3-V. Hair 5-IV usually double (2,3b).

Paddle: Index usually 1.4-1.6 (1.3-1.7).

LARVA (fig. 20). Head: 1.12 mm. Siphon: 1.17 mm. Anal Saddle: 0.41 mm.

Head: Hair 5-C usually double (1-3b). Hair 6-C 3,4b (2-4). Hair 14-C usually dou-

ble. Abdomen: Hair 1-II-V usually 3,4b (2-5), the branches relatively stout and

coarse to near apex. Hair 2-II usually 2-4b. Hair 5-II-[V usually 3,4b (2-4) and

5-V-VII usually single or double (1-3b), the branches relatively stout and coarse

to near apex. Hair 13-II-V usually 3-5b (3-6), the branches relatively stout and

coarse to near apex; 13-VI usually 8-15b (7-17), the branches much shorter and

finer than those of 13-VII; hair 13-VII usually 3,4b. Segment VIII: Branches of

hair 1 subequal in strength to or stronger than branches of 1-X. Comb scales us-

ually 25-42 (24-49); apex of larger scales usually fringed. Siphon: Index about

3.9-4.6 (3.5-5.0). Anal Segment: Hair 4a-X long, usually at least 2.0 length of

anal saddle.

SYSTEMATICS. Aedes schicki is readily separated from the most closely re-

lated species, chionotum, as follows: in the adult by (1) the shorter and broader

erect scales on the head of the female, (2) the shorter palpus of the male, (3)

the presence of scales on the metameron, and (4) the conspicuous patch of white

scales at the base of hindtarsal segment 1; and in the larva by hairs 1-I-VI, 5-

II-VII and 13-II-V, which have fewer, rigid, stout branches. Pupae of the 2 spec-

ies are apparently indistinguishable. The presence of a definite patch of scales

on the metameron but not on the prosternum of the adult and the stout rigid

hairs of the larva of schicki are unique developments within the Kompi Group.

BIONOMICS. The immatures of schicki are found in treeholes where they may

be associated with the immatures of Aedes (Ochlerotatus) muelleri. The habits

of the adults are unknown. The species is apparently restricted to montane oak-

pine forest.

DISTRIBUTION (fig. 10). Aedes schicki has been collected at elevations from

2350 to 2620 meters in the Sierra Madre Occidental in northcentral Mexico. Ma-

terial examined: 83 specimens; 16 males, 12 females, 29 larvae, 26 pupae; 24

individual rearings (20 larval, 4 pupal).

MEXICO. Durango: Durango (52 km W), 3 July 1967, G.A. Schroeder (MEX 488), 3 IpF

(488-11-13), 2 P, 6 L [UCLA]. La Emerta (4 km E), type series, see above.

8. Aedes (Protomacleaya) chionotum Zavortink, n.sp.

Figs. 10,13,21,22

TYPES: Holotype female with associated larval and pupal skins (MEX 304-20), 6.9 km N

of Cuernavaca, Morelos, Mexico, elevation near 2080 m, larva from treehole, 19 Aug 1965,

48 Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

R.X. Schick and D.A. Schroeder [USNM]. Allotype male with associated pupal skin and geni-

talia (MEX 304-11), same data as holotype [USNM]. Paratypes: 2 IpF (304-19,22), 2 pM (304-

10,12), 6 Ip (304-15-18,21,23), 2 M gen, 1 P, 5 L, same data as holotype (MEX 304) [UCLA,

ISET, BM].

FEMALE (fig. 13). Wing: 4.17 mm. Proboscis: 2.39 mm. Forefemur: 2.20 mm.

Abdomen: about 2.7 mm. Head: Erect scales long and narrow, their width at

apex scarcely greater than width at midheight. Thorax: Anterior dorsocentral bris-

tles numerous and well developed; humeral, lateral prescutal and posterior fossal

bristles few. Dark scaled areas of mesoscutum with numerous scattered yellowish

or whitish scales. Light mesoscutal lines relatively inconspicuous; acrostichal line

absent to narrow and broken; lateral prescutellar line narrow and broken to mod-

erately broad; posterior outer dorsocentral line narrow; lateral marginal line nar-

row; posterior fossal line narrow, sometimes broadened and denser mesally and

then appearing as a patch. Midlobe of scutellum entirely or predominantly light

scaled. Most pleural patches moderately dense and moderately well defined, con-

sisting of broad flat to outstanding disheveled white scales. Upper anterior ppn

with scales sparse, most narrow, curved; pst and metameron bare. Legs: Posterior

surface of hindfemur and midfemur with white streak extending from base to or

near apex. Foretibia and midtibia frequently streaked with white from near base

to near apex on posterior surface. Foretarsal segment 1 usually marked with white

on ventral and posterior surfaces in at least basal 0.4; midtarsal segment 1 usually

marked with white on dorsal and posterior surfaces in at least basal 0.5; hind-

tarsal segment 1 sometimes with a few light scales at base of dorsal surface.

MALE. Similar to female except for sexual characters and mesoscutal ornamen-

tation. Head: Erect scales not unusually long or narrow. Palpus subequal in length

to or longer than proboscis; with numerous long bristles from apex of segment

3 distad. Thorax: Mesoscutum entirely silver-white scaled or with a few pos-

terior inner dorsocentral scales dark. Pleural patches with scales less disheveled

than in female. Upper anterior ppn with scales denser, broader and flatter than

in female. Wing: Remigium (base of vein R) partially to entirely light scaled. Ab-

domen: Tergites with conspicuous white scaled basal band.

MALE GENITALIA. (fig. 21). Apparently indistinguishable from other species

in group.

PUPA (fig. 21). Abdomen: about 3.7 mm. Trumpet: 0.73 mm. Paddle: 0.95

mm. Apparently indistinguishable from schicki. Cephalothorax: Hair 10-C strongly

developed, subequal in stoutness to and longer than 12-C. Abdomen: Float hair

(1-I) strongly developed, with longest branches not extending to lateral margin

of segment I, and with primary branches with or without secondary branches;

branches of 1-II usually 6-8 (5-8), strongly developed, as stout as primary branches

of 1-I; hair 1-III-V 3-6b. Hair 2-IV-VI usually in line with or laterad of hair 1

of corresponding segment and usually relatively far from caudal margin of seg-

ment; 2-V far cephalad of level of 3-V. Hair 5-IV usually double (1-4b). Paddle:

Index 1.5-1.6.

LARVA (fig. 22). Head: 1.04 mm. Siphon: 1.11 mm. Anal Saddle: 0.40 mm.

Head: Hair 5-C usually 3b (2-4). Hair 6-C usually 4,5b (4-7). Hair 14-C usually

2-4b (1-5). Abdomen: Hair 1-II-V usually 5-8b (5-10), the branches normal. Hair

2-II usually 4-6b (3-8). Hair 5-II-[V usually 6-8b (5-10) and 5-V-VII usually 4,5b

(3-6), the branches normal. Hair 13-II-V usually 6-9b (4-11), the branches nor-

mal; 13-VI usually 9-14b (7-17), the branches much shorter and finer than those

of 13-VII; hair 13-VII usually 5,6b (4-7). Segment VIII: Branches of hair 1 sub-

Zavortink: New World Aedes 49

equal in strength to or finer than branches of 1-X. Comb scales usually 24-34

(21-40); apex of larger scales usually fringed. Siphon: Index about 3.5-4.1. Anal

Segment: Hair 4a-X long, usually at least 2.0 length of anal saddle.

SYSTEMATICS. Aedes chionotum is closely related to schicki and may be dis-

tinguished from it as follows: in the adult by (1) the long narrow erect scales

on the head of the female, (2) the longer palpus of the male, (3) the absence

of scales on the metameron, and (4) the absence of a conspicuous patch of white

scales at the base of hindtarsal segment 1; and in the larva by hairs 1-I-VI, 5-

II-VII and 13-II-V, which have more numerous, flexible, tapering branches. The

pupae of the 2 species appear to be indistinguishable. The long narrow erect

scales on the head of the female and the highly branched larval hairs serve to

separate chionotum from all other species of the Kompi Group.

BIONOMICS. The immature stages of chionotum have been found in treeholes;

they have not been associated with any other species of mosquito. Habits of the

adults are unknown. This species is apparently restricted to high elevation forests.

DISTRIBUTION (fig. 10). Aedes chionotum is known from the Mexican States

of Morelos and Oaxaca at elevations of 2080 to 2210 meters. Material examined:

74 specimens; 3 males, 3 females, 55 larvae, 13 pupae; 12 individual rearings (3

larval, 3 pupal, 6 incomplete).

MEXICO. Morelos: Cuernavaca (6.9 km N), type series, cited above. Oaxaca: Vivero Rancho

Teja, Ixtlan de Juarez, 3 July 1970, D.A. and K. Schroeder (MEX 518), 35 L; same data (MEX

521), 6 L [UCLA].

9. Aedes (Protomacleaya) niveoscutum Zavortink, n.sp.

Figs. 10,13,23,24

TYPES: Holotype male with associated larval and pupal skins (MEX 669-12), El Mirador

(2.4 km S Quililla), Jalisco, Mexico, elevation about 1710 m, larva from rothole in living oak

tree, 8 June 1971, T.J. Zavortink and L.T. Nielsen [USNM]. Allotype female with associated

larval and pupal skins (MEX 675-18), same data as holotype [USNM]. Paratypes: 1 lpM (669-

14), 2 lpF (669-11,16), 1 M, 1 P, same data as holotype (MEX 669); 2 IpM (670-10,11), 1 M

gen, same data as holotype (MEX 670); 2 IpF (671-10,11), 1 M, 1 P, same data as holotype

(MEX 671); 1 IpM (675-10), 3 lpF (675-20,21,24), 61 M, 3 M gen, 22 F, 83 P, 18 L, same

data as holotype (MEX 675) [UCLA, BM, ISET]; 2 lpM (1,2), 1 lpF (4), 1 pM (8), same data

as holotype (N-22-71); 1 IpM (3), same data as holotype (N-23-71); 1 lpM (8), 2 lpF (5,6),

same data as holotype (N-27-71) [UTAH].

FEMALE (fig. 13). Wing: 4.21 mm. Proboscis: 2.73 mm. Forefemur: 2.55 mm.

Abdomen: about 3.1 mm. Head: Erect scales moderately long and moderately

broad, their width at apex 2 or 3 times greater than width at midheight. Thorax:

Anterior dorsocentral bristles numerous and well developed; humeral, lateral pre-

scutal and posterior fossal bristles few. Dark scaled areas of mesoscutum without

yellowish or whitish scales. Light mesoscutal lines predominantly inconspicuous;

acrostichal line complete, narrow to moderately broad; lateral prescutellar line us-

ually narrow and broken; posterior outer dorsocentral line poorly developed, nar-

row and incomplete; lateral marginal line usually moderately broad anteriorly, re-

mainder narrow to moderately broad; posterior fossal line absent to weakly de-

veloped and narrow, never broadened mesally to form a patch. Midlobe of scu-

tellum with scales predominantly dark. Most pleural patches dense and well de-

fined, consisting of broad flat to appressed imbricate scales. Upper anterior ppn

50 Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

with scales usually dense, moderately broad to broad, appressed; pst with well

developed oblique patch of scales; metameron bare or with a few scales. Legs:

Posterior surface of forefemur and midfemur with white streak extending from

base to or near apex. Foretibia and usually midtibia streaked or speckled with

white on posterior surface. Foretarsal segment 1 usually marked with white on

at least posterior and ventral surfaces in at least basal 0.4; midtarsal segment |

usually marked with white on at least dorsal, posterior and ventral surfaces in

at least basal 0.4; hindtarsal segment 1 dark scaled.

MALE. Similar to female except for sexual characters and mesoscutal ornamen-

tation. Head: Palpus shorter than proboscis; with relatively few long bristles from

apex of segment 3 distad. Thorax: Mesoscutum entirely silver-white scaled or rarely

with a few posterior inner dorsocentral scales dark. Wing: Remigium (base of vein

R) entirely dark scaled. Abdomen: Tergites without basal band.

MALE GENITALIA (fig. 23). Apparently indistinguishable from other species

of group.

PUPA (fig. 23). Abdomen: about 3.2 mm. Trumpet: 0.58 mm. Paddle: 0.83

mm. Apparently indistinguishable from sandrae. Cephalothorax: Hair 10-C mod-

erately to strongly developed, subequal in stoutness and length to 12-C or stouter

and longer. Abdomen: Float hair (1-1) moderately to strongly developed, with

longest branches sometimes extending to lateral margin of segment I, and with

primary branches with or without secondary branches; branches of 1-II usually

4-6 (3-7), moderately to strongly developed and finer than or subequal in stout-

ness to primary branches of 1-I; hair 1-III-V usually 2-4b (2-5). Hair 2-[V-VI us-

ually in line with or laterad of hair 1 of corresponding segment and usually mod-

erately far from caudal margin of segment; 2-V usually slightly to greatly ceph-

alad of level of 3-V. Hair 5-IV usually single (single or double). Paddle: Index

usually 1.2-1.3 (1.2-1.4).

LARVA (fig. 24). Head: 1.07 mm. Siphon: 0.92 mm. Anal Saddle: 0.40 mm.

Head: Hair 5-C usually double (1-3b). Hair 6-C usually single or double (1-3b).

Hair 14-C usually double (single or double). Abdomen: Hair 1-II-V usually 3,4b

(2-5), the branches normal. Hair 2-I] usually 2,3b (2-4). Hair 5-H-VI usually 3-5b

(3-6) and 5-VII usually 2,3b, the branches normal. Hair 13-II-V usually 3-5b (2-

6), the branches normal; 13-VI usually 4-6b (4-8), the branches shorter and us-

ually slightly finer than those of 13-VII; hair 13-VII 3,4b. Segment VIL: Branches

of hair 1 finer to stouter than branches of 1-X. Comb scales usually 25-40 (22-

50); apex of larger scales evenly fringed or 1 or 2 apical elements of fringe slightly

enlarged. Siphon: Index usually 2.7-3.0 (2.5-3.1). Anal Segment: Hair 4a-X long,

usually at least 2.0 length of anal saddle.

SYSTEMATICS. This species differs from the closely related sandrae as follows:

in the adult by (1) the shorter and sparsely bristled palpus of the male, (2) the

absence of a white scaled patch at the union of the posterior fossal and posterior

outer dorsocentral lines on the mesoscutum of the female, (3) the presence of

a patch of scales on the prosternum, and (4) the dark scaled remigium (base of

vein R) of the wing of the male; and in the larva by (1) the absence of a definite

moderate-sized spine at the apex of the larger comb scales, and (2) the longer

hair 4a-X. Pupae of niveoscutum and sandrae are apparently indistinguishable.

Aedes niveoscutum differs from all other members of the Kompi Group in the

presence of a well developed patch of scales on the prosternum but not on the

metameron of the adults and in the sparsely bristled palpus of the male.

There is considerable variation in the single population of niveoscutum that

Zavortink: New World Aedes 51

has been studied. In the adults the normally short palpus of the male is occa-

sionally as long as the proboscis, the usually narrow lateral prescutal line of the

female is sometimes broadened, and the typically bare metameron often has a

few scales. Since all of these variations are in the direction of burgeri, which oc-

curs with this population of niveoscutum, they are probably the result of hybrid-

ization with that species. In the immatures the branches of many hairs are fre-

quently reduced in number and strength. This variability may also be due to hy-

bridization with burgeri or, since variability of this type and degree is exhibited

by the immatures of the closely related sandrae, it may be inherent in this lineage.

BIONOMICS. The immature stages of niveoscutum are found in treeholes where

they may be associated with burgeri. Habits of the adults are unknown.

DISTRIBUTION (fig. 10). This species is presently known from only the type

locality in the State of Jalisco, Mexico, at an elevation of about 1710 meters.

It probably extends southward at appropriate elevations in the Sierra Madre del

Sur. Material examined: 251 specimens; 74 males, 33 females, 38 larvae, 106 pu-

pae; 21 individual rearings (20 larval, 1 pupal).

MEXICO. Jalisco: El Mirador, type series, cited above.

10. Aedes (Protomacleaya) sandrae Zavortink, n.sp.

Figs. 10,13,25,26

TYPES: Holotype female with associated larval and pupal skins (GUA 125-11), Hipodromo

del Norte in Guatemala City, Guatemala, Guatemala, elevation near 1500 m, larva from cut or

broken bamboo internode, 4 Sept 1964, V.P. Cowsill and W. Almengor [USNM]. Allotype male

with associated pupal skin and genitalia (GUA 125-100), same data as holotype [USNM]. Para-

types: 1 \IpF (125-12), 2 pF (125-103,105) (GUA 125), same data as holotype [UCLA]. This

species is dedicated to Sandra J. Heinemann in recognition of her contributions to the project

“Mosquitoes of Middle America.”

FEMALE (fig. 13). Wing: 4.13 mm. Proboscis: 2.64 mm. Forefemur: 2.31 mm.

Abdomen: about 3.0 mm. Head: Erect scales moderately long and moderately

broad, their width at apex 2-4 times greater than width at midheight. Thorax:

Anterior dorsocentral bristles numerous and well developed; humeral, lateral pre-

scutal and posterior fossal bristles few. Dark scaled areas of mesoscutum without

scattered yellowish or whitish scales. Light mesoscutal lines conspicuous; acros-

tichal and posterior outer dorsocentral lines narrow; lateral prescutellar line nar-

row, frequently broken; lateral marginal line broadened anteriorly, remainder nar-

row to moderately broad; posterior fossal line broadened mesally, forming a con-

spicuous patch. Midlobe of scutellum predominantly dark scaled. Most pleural

patches very dense and well defined, consisting of broad closely appressed im-

bricate silver-white scales. Upper anterior ppn with scales usually dense, moder-

ately broad to broad, appressed and imbricate; pst and metameron bare. Legs:

Posterior surface of forefemur and midfemur with white streak of basal portion

narrowing and extending to or near apex. Foretibia and midtibia usually weakly

marked with white on posterior surface. Foretarsal segment | marked with white

on ventral and posterior surfaces in at least basal 0.3; midtarsal segment | marked

with white on all surfaces in at least basal 0.3; hindtarsal segment 1 dark scaled.

MALE. Essentially as in female except for sexual characters and mesoscutal

ornamentation. Head: Palpus subequal in length to proboscis; apparently with

52 Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

numerous long bristles from apex of segment 3 distad. Thorax: Mesoscutum en-

tirely silver-white scaled. Wing: Remigium (base of vein R) entirely light scaled.

Abdomen: Some tergites without basal band, some with narrow light scaled band.

MALE GENITALIA (fig. 25). Apparently indistinguishable from other species

in group.

PUPA (fig. 25). Abdomen: about 3.4 mm. Trumpet: 0.67 mm. Paddle: 0.82.

mm. Apparently indistinguishable from niveoscutum. Cephalothorax: Hair 10-C

moderately to strongly developed, usually slightly stouter and longer than 12-C.

Abdomen: Float hair (1-I) very strongly developed, with longest branches extend-

ing at least to lateral margin of segment I, and with primary branches with or

without secondary branches; branches of 1-II usually 4-6 (3-7), strongly devel-

oped and subequal in stoutness to primary branches of 1-I; hair 1-III-V usually

3-5b (1-5). Hair 2-IV-VI usually in line with or laterad of hair 1 of correspond-

ing segment and usually relatively far from caudal margin of segment; 2-V far

cephalad of level of 3-V. Hair 5-IV single. Paddle: Index 1.1-1.3.

LARVA (fig. 26). Head: 0.92 mm. Siphon: 0.83 mm. Anal Saddle: 0.32 mm.

Head: Hair 5-C usually 2,3b (1-4). Hair 6-C usually 2-4b (1-5). Hair 14-C usu-

ally double. Abdomen: Hair 1-II-V usually 2-4b (1-6), the branches normal. Hair

2-II usually 2,3b (2-4). Hair 5-II-VII 3-5b, the branches normal. Hair 13-II-V us-

ually 3-5b (2-6), the branches normal; 13-VI usually 4b (3-5), the branches sim-

ilar in length and stoutness to those of 13-VII; hair 13-VII 3,4b. Segment VIII:

Branches of hair 1 subequal in strength to or finer or stouter than branches of

1-X. Comb scales usually 24-33 (22-38); apex of larger scales produced into a

moderate-sized spine. Siphon: Index about 2.7-3.9. Anal Segment: Hair 4a-X short,

usually only 1.0-1.5 length of anal saddle.

SYSTEMATICS. Aedes sandrae is most- closely related to niveoscutum from

which it differs as follows: in the adult by (1) the longer and densely bristled

palpus of the male, (2) the presence of a conspicuous white scaled patch at the

juncture of the posterior fossal and posterior outer dorsocentral lines on the meso-

scutum of the female, (3) the absence of a patch of scales on the prosternum,

and (4) the presence of light scales on the remigium (base of vein R) of the

wing in the male; and in the larva by (1) the presence of a definite moderate-

sized spine at the apex of the larger comb scales, and (2) the shortened hair

4a-X. Pupae of the 2 species appear to be indistinguishable. Aedes sandrae is dif-

ferentiated from all other species in the Kompi Group by the 2 larval characters

mentioned above.

There is considerable variability in the number-and strength of the branches

of many hairs in the larva and pupa. Particularly noticeable is the variation in

hairs 5,6-C on the head of the larva. This variation is frequently, but not invar-

iably, correlated with dimorphism of the antennae and mouthbrushes, as follows:

larvae with hairs 5,6-C single or double usually have short antennae and pectinate

mouthbrushes; those with hairs 5,6-C 3-5-branched usually have long antennae and

simple mouthbrushes.

BIONOMICS. Larvae and pupae of sandrae have been collected in treeholes

and broken or cut bamboo internodes. At lower elevations they have been assoc-

iated with Aedes (Protomacleaya) podographicus Dyar and Knab, 1906, a mem-

ber of the Terrens Group. The single female from Mexico was taken in. a biting-

landing collection with human bait at dusk; adults of Ae. (P.) sumidero Schick,

1970, another species of the Terrens Group, were associated with it.

DISTRIBUTION (fig. 10). Aedes sandrae has been found in southern Mexico

Zavortink: New World Aedes 53

and western Guatemala at elevations of 200 to 2000 meters. Material examined:

35 specimens; 1 male, 5 females, 24 larvae, 5 pupae; 5 individual rearings (2 lar-

val, 3 pupal).

GUATEMALA. Esquintla: Esquintla (7-8 km SW), 10 July 1964, V.P. Cowsill (GUA 41),

1 L [UCLA], Guatemala: Guatemala City, Bethania, 15 July 1964, T.J. Zavortink and V.P.

Cowsill (GUA 44), 2 L [UCLA]. Guatemala City, Hipodromo del Norte, type series, see above.

Solola: San Andres Semetabal, 17 July 1964, V.P. Cowsill (GUA 51), 19 L [UCLA].

MEXICO. Chiapas: Sumidero (24 km N Tuxtla Gutierrez), 17 Aug 1964, E. Fisher and D.

Verity (MEX 128), 1 F [UCLA].

KNABI GROUP

11. Aedes (Protomacleaya) knabi (Coquillett)

Figs. 10,27,28

1906. Culex knabi Coquillett, 1906b:183-184. TYPE: Lectotype female with associated pupal

skin (291a), Tehuantepec, Oaxaca, Mexico, 1 July 1905, F. Knab [USNM; selection

of Stone and Knight, 1956:220].

Aedes (Finlaya) knabi of Dyar (1928:226-227); Edwards (1932:152); Knight and Marks (1952:

538,563); Stone, Knight and Starcke (1959:164).

Aedes knabi of Dyar (1906:16); Dyar and Knab (1906:203); Howard, Dyar and Knab (1917:

841-842).

Aedes (?Gualteria) knabi of Dyar (1918:73).

Aedes (Ochlerotatus) knabi.of Dyar (1922d:160).

Ochlerotatus knabi of Coquillett (1906c: 18,21).

FEMALE. Wing: 5.14 mm. Proboscis: 2.96 mm. Forefemur: 3.13 mm. Abdo-

men: about 3.7 mm. Head: Integument yellow to tan. Erect scales and narrow

curved scales unicolorous, golden. Thorax: Integument yellow to tan. Acrostichal,

anterior dorsocentral, humeral, lateral prescutal and posterior fossal bristles few,

moderately to strongly developed. Mesoscutum entirely covered with small yel-

low or golden scales and larger cream-yellow, yellow or golden scales except for

inconspicuous to ‘conspicuous inner dorsocentral line of small bronzy or brown

scales. Midlobe of scutellum with scales sparse or dense, narrow and curved or

broad and flat, yellow to golden. Ppn scales broad and flat and/or narrow and

curved, yellow to golden. Pleural scale patches silver-white; psp scales absent; ssp

scales absent or few; pst and metameron bare. Legs: Posterior or ventral surface

of forefemur and midfemur with light scales restricted to basal 0.5-0.7. Knee spot

absent on foreleg, absent to small on midleg, small on hindleg. Tibiae dark scaled

or with short pale streak at base of posterior surface on foreleg and midleg. Tar-

sal segment 1 of foreleg and midleg with poorly defined light ring in basal 0.5-

1.0; tarsal segment 1 of hindleg with very poorly defined light streak or band

in basal portion; tarsal segment 2 of all legs with light ring or band in basal 0.4-

1.0; tarsal segments 3 and 4 of foreleg and midleg sometimes light scaled dor-

sally. Claws of hindleg with acute submedian tooth. Wing: Without light scales

at base of costa or vein R. Abdomen: Tergites II-VIII dark scaled with baso-

lateral light patch. Basal sternites light scaled, distal dark scaled with basolateral

light patch or basal light band that is broader laterally.

MALE. Essentially as in female except for sexual characters. Head: Palpus en-

54 Contrib. Amer. Ent. Inst., vol. 8, no. 3, 1972

tirely dark scaled; palpal bristles not discernible. Thorax: Mesoscutum entirely

golden scaled except for a very few posterior inner dorsocentral brown scales.

Legs: Larger claw of foreleg and midleg with very small basal external tooth;

claws of hindleg with an acute submedian tooth. Wing: Remigium (base of vein

R) dark scaled.

MALE GENITALIA (fig. 27). Sidepiece: Apparently weakly to moderately pig-

mented. Basal tergomesal area slightly swollen; setae of basal tergomesal area rel-

atively numerous, apparently arising from a moderately sclerotized plaque; med-

ian sternomesal sclerite moderately developed; median sternomesal tuft apparently

moderately developed, the setae apparently strongly curved dorsad. Claspette: Fil-

ament angulate. Phallosome: Aedeagus relatively long and slender, length about

2.5 greatest width.

PUPA (fig. 27). Abdomen: about 4.0 mm. Trumpet: 0.60 mm. Paddle: 1.05

mm. Hairs moderately strong and moderately pigmented. Cephalothorax: Hairs

4,8-C weakly to moderately developed, short; 8-C shorter and finer than 9-C.

Trumpet: Golden brown to brown. Abdomen: Hair 1-II with 4-6 moderately de-

veloped short branches; 1-III-V double. Hair 6-I,]] moderately developed, much

finer than 3-I,II. Hair 9-III-VI fine, short. Paddle: Apex produced.

LARVA (fig. 28). Head: 1.25 mm. Siphon: 1.32 mm. Anal Saddle: 0.54 mm.

Head: Labrum produced medially in dorsal aspect. Hair 1-C fine, arising relatively

far from edge of labrum. Hair 4-C moderately developed, with 6-10 moderately

long branches; located midway between midline and 6-C. Hairs 5,6-C single. Hair

7-C 2-4b. Hair 15-C enlarged, 25-35b. Mental plate not distinctly triangular; 1

or 2 of the more lateral teeth conspicuously enlarged. Antenna: Shaft with num-

erous small spicules. Thorax: Pigment absent. Hairs 1,5-P single. Abdomen: Hair

12-I present. Segment VIII: Comb scales 16-30, in an irregular patch. Siphon:

Index about 3.0. Acus attached. Pecten: teeth 28-47. Hair 1-S single. Anal Seg-

ment: Saddle large, extending 0.75-0.85 down lateral surface of segment; appar-

ently without a narrow lightened or transparent ventral margin or submarginal

area. Hair 1-X single. Ventral brush with 7 pairs of hairs; hairs 4b,4c-X 5-7b.

Gills tapered distally; dorsal subequal to ventral in length, about 1.0-1.2 length

of saddle.

SYSTEMATICS. Aedes knabi is characterized as follows: in the adult by (1)

the yellow to tan integument of the head and thorax, (2) the predominantly yel-

low to golden scales of the head and mesoscutum, and (3) the toothed claws of

the hindleg; in the male genitalia by the differentially sclerotized plaque from

which the setae of the basal tergomesal area arise; in the pupa by the combina-

tion of (1) the short hairs 4,8-C, and (2) the pointed or produced paddle apex;

and in the larva by (1) the produced labrum, (2) the fine hair 1-C, (3) the en-

larged and many branched 15-C, and (4) the much enlarged lateral teeth of the

mental plate.

Aedes knabi has so many unique developments, especially in the adult and larva,

that I am placing it into a separate group. The many unusual characteristics of

this species indicate a long history for the lineage leading to it.

Aedes knabi is known from only the small type series and 1 male with its

associated pupal skin and 1 larva in the UCLA collection. All of the immatures

and the only male genitalia are in very poor condition and inaccuracies may have

been introduced into the illustrations reconstructed from these.

BIONOMICS. The immatures of knabi have been collected only in treeholes.

They were associated with Ae. (P.) tehuantepec Schick, 1970 in 1 collection. All

Zavortink: New World Aedes 55

known adults are reared and nothing is known about their habits.

DISTRIBUTION (fig. 10). Aedes knabi is known at present from only the vicin-

ity of Tehuantepec, Oaxaca, Mexico, at elevations below 100 meters. Material ex-

amined: 14 specimens; 1 male, 5 females, 4 larvae, 4 pupae; 4 individual rearings

(2 larval, 1 pupal, 1 incomplete).

MEXICO. Oaxaca: Salina Cruz, 15 July 1905, F. Knab (307), 2 IpF (307d,g), 1 lp (307c),

2 F (307a,b) [USNM]; 9 Aug 1964, D. Verity (MEX 104), 1 pM (104-10), 1 M gen, 1 L

[UCLA]. Tehuantepec, 1 July 1905, F, Knab (291), 1 F (291b) [USNM].

Subgenus KOMPIA Aitken

1941. Kompia Aitken, 1941:81-82. TYPE SPECIES: Aedes (Kompia) purpureipes Aitken,

1941, Mexico; monobasic and original designation.

12. Aedes (Kompia) purpureipes Aitken

Figs. 29-32

1941. Aedes (Kompia) purpureipes Aitken, 1941:82-84. TYPE: Holotype female, Triunfo,

Baja California Sur, Mexico, 7 July 1938, A.E. Michelbacher and E.S. Ross [CAS].

Aedes (Kompia) purpureipes of Aitken (1942:167); Belkin and McDonald (1957:190); McDonald

(1957b:529-535); Mattingly (1961:38); Stone (1967:208); Zavortink (1970:8).

Aedes (Ochlerotatus) purpureipes of Vargas (1949:261-265; 1950:64-65); Carpenter and LaCasse

(1955:220-221); Mattingly (1958:6); Stone, Knight and Starcke (1959:151).

FEMALE (fig. 30). Wing: 3.13 mm. Proboscis: 2.29 mm. Forefemur: 2.02 mm.

Abdomen: about 2.9 mm. Dark scales of head, proboscis, palpus, legs and abdo-

men with metallic blue, violet or purple reflections. Head: Eyes broadly separ-

ated above antennae, the resulting space with broad flat silver scales. Integument

light to dark brown. Frontal bristles absent. Orbital bristles relatively few, mesal

4-7 pairs strongly developed, lateral 3 or 4 pairs moderately developed. Vertex

with relatively few black proclinate erect scales and with broad flat silver scales

and sometimes appressed curved creamish-yellow to golden scales medially, broad

flat black scales submedially and broad flat silver scales laterally. Occiput with

numerous upright black and yellow erect scales and creamish-yellow to golden

predominantly narrow and curved decumbent scales. Orbital line of broad flat sil-

ver scales present. Dorsolateral surface with black, silver and yellowish broad

flat scales. Lateral and ventral surfaces with yellowish broad flat scales. Clypeus

moderately large, bare. Proboscis relatively stout, longer than forefemur; entirely

dark scaled; with a few basal bristles. Palpus moderately long, about 0.24-0.27

length of proboscis: 4-segmented; segment 4 moderate in size; segments 1-3 with

bristles; entirely dark scaled. Antenna slightly shorter than proboscis; torus with

conspicuous large patch of broad flat silver scales; flagellar segment 1 longer

than 2, usually slightly swollen, with small dark scales; flagellar segments 2-13

usually with 6 moderate bristles in basal whorl; each of flagellar segments 3-6

or 7 slightly longer than preceding segment, segments 6 or 7-12 subequal. Tho-

rax: Integument yellow to golden with light to dark brown areas. Acrostichal

bristles usually restricted to anterior promontory; dorsocentral bristles present

caudad to level of ppn bristles; prescutellar and supraalar bristles numerous and