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Contributions
of the
American Entomological Institute
Volume 5, Number 1, 1969
i
MOSQUITO STUDIES (Diptera, Culicidae)
A new species of treehole breeding Aedes (Ochlerotatus)
from southern California. By Thomas.J. Zavortink.
Two new species of Deinocerites from Costa Rica.
By Abdiel J. Adames and Charles L. Hogue.
The subgenus Micraedes of Culex. By O. G. W. Berlin.
a HSOlV l4 yy
MAR 23 19/0
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CONTRIBUTIONS
of the
AMERICAN ENTOMOLOGICAL INSTITUTE
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(Continued on back cover)
MOSQUITO STUDIES (Diptera, Culicidae)
XVI. A NEW SPECIES OF TREEHOLE BREEDING AEDES (OCHLEROTATUS)
FROM SOUTHERN CALIFORNIA’
By
Thomas A # Zavortink”
In April 1964 a large number of Aedes larvae were taken from a rot hole in a syc-
amore tree growing near the junction of Santiago and Little Rock creeks, on the
north (desert) slope of the San Gabriel Mountains, Los Angeles County, California.
Very few of these larvae were reared to adulthood; the majority, most of them
younger instars, were killed and preserved. This material was not examined until
January 1969, at which time it was discovered that 2 different species were present.
One of these was the common Pacific Coast treehole mosquito, Aedes sierrensis
_ (Ludlow, 1905), the second appeared to be an undescribed species belonging, as does
sierrensis, to the varipalpus complex of the subgenus Ochlerotatus. Additional mater-
ial collected during February and March 1969 indicated that this second species was
distinct from the 2 remaining species of the varipalpus complex, monticola Belkin &
McDonald, 1957 and varipalpus (Coquillett, 1902), and that it was widespread in
the desert drainages of southern California. This new species, deserticola, is described
and illustrated in the present paper.
I wish to thank John N. Belkin, George W. Berlin and Robert X. Schick for read- .
ing the manuscript and offering valuable suggestions. I am indebted to the following
individuals for help in the collecting, rearing and preparation of specimens, for ink-
ing the plates and typing the manuscript: Caryle Abrams, Judith A. Bergland, Sheila
_E. Bernstein, Dennis W. Heinemann, Sandra J. Heinemann, Nancy L. Martsch and
William A. Powder.
‘Aedes (Ochlerotatus) deserticola Zavortink, n.sp.
Figs, 1.2
TYPES: Holotype 6 with associated larval and pupal skins (UCLA 516-32), junction of Santi-
ago and Little Rock creeks, about 5 air mi southwest of Littlerock, San Gabriel Mountains, Los
Angeles County, California, larva from a rot hole in a living cottonwood tree, 4 Feb 1969, T.J.
Zavortink [USNM]. Allotype ° with associated larval and pupal skins (UCLA 516-33), same data
‘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.
2 Contrib. Amer. Ent. Inst., vol. 5, no. 1, 1969 ©
as holotype [USNM] . Paratypes: 2 Ipd (516-10,34), 17 Ip? (S516-35-51), 1 lp (516-31), 17 6, 40 P,
78 L, same data as holotype (UCLA 516); 5 Ipd (517-30-34), 18 lp? ($17-35-39,41-53), 1 lp (S17-
40), 19 6, 17 9, 39 P, 125 L, same data as holotype except collected in a sycamore treehole
(UCLA 517); 4 Ipd (522-10-13), 4 Ip? (522-15-17,19), 11 6,4 9, 13 P, 8 L, same data as holotype
except collected in a sycamore treehole (UCLA 522); 3 6,5 9, 146 L, same data as holotype ex-
cept collected in a sycamore treehole on 4 Apr 1964 (UCLA 503) [UCLA].
FEMALE. Wing: 3.70 mm. Proboscis: 2.14 mm. Forefemur: 1.82 mm. Abdomen: _
about 2.9 mm. Very similar to sierrensis and other members of group, differing pri-
marily in the following. Head: Proboscis entirely dark scaled or with a small patch
of light scales at base on ventral surface. Thorax: Mesonotum with broad median
stripe of pale golden scales; at least some of the light scales immediately laterad of
posterior dorsocentral and lateral prescutellar bristles yellowish. Pleural scale patches
relatively small and pleural bristles relatively few; ssp and lower mep bristles absent;
postcoxal area and metameron without scales. Legs: Hindtarsal segment 4 entirely
dark scaled or with a few white scales at ends.
MALE. Similar to female except for usual sexual differences. Labium: Basal half
light scaled on at least ventral surface. Palpus: Without a patch of light scales over
the joint between segments 2 and 3.
MALE GENITALIA (fig. 1). As figured; very similar to monticola and varipalpus.
Segment IX: Tergite lobe usually with 4-6 (2-7) strong bristles. Sidepiece: Basal ter-
gomesal lobe with 1 strongly differentiated seta and. 1-3 long slender bristles; tergo-
mesal margin without hairs. Claspette: Connection between claspettes declivous mes-
sally. Clasper: Strongly curved apically. Spiniform: Relatively long.
PUPA (fig. 1). Abdomen: 3.12 mm. Trumpet: 0.62 mm. Paddle: 0.79 mm. Posi-
tion, length, degree of development and modal number of branches of all hairs as
figured. Very similar to other members of group, distinguished on the basis of the
following characters. Cephalothorax: Integument more or less uniformly light yel-
lowish in color. Trumpet: Bright yellowish brown in color. Abdomen: Integument
more or less uniformly light yellowish in color; hair 1-II-VII subequal in develop-
ment, usually double (1-4b) on segment II, usually single (single, double) on seg-
ments ITI-VII; 5-[V-VI usually relatively long and exceeding 9-VII in length. Paddle:
Integument uniformly light yellowish except for darker midrib; basal portion of ex-
ternal buttress often straight or slightly concave.
FOURTH INSTAR LARVA (fig. 2). Head: 0.81 mm. Siphon: 0.69 mm. Anal Sad-
dle: 0.20 mm. Position, length, degree of development and modal number of branch-
es of all hairs as figured. Similar to other species in the complex, especially monticola
and varipalpus, but distinguished as follows. Head: Integument light yellowish brown
in color with ocular area lighter and posterior portion darker; sculpturing indistinct.
Antenna: Uniformly light yellowish brown. Thorax: Epidermis and fat body with-
out pigment, living larva white; integument without spicules. Abdomen: Pigmenta-
tion and spiculation as for thorax; hair 7-I JI usually weakly developed, similar to
hair 10 of corresponding segment; 1,13-IJJ-V usually double, with outer branch
much longer and stronger than inner and usually longer than siphon. Segment VIII:
Comb scales usually brown and 7-12 (5-15) in number, arranged in a single regular or
irregular row; hairs 1,3 frequently single. Siphon: Index usually 2.2-2.4 (2.1-2.6); pig-
mentation uniformly brown; sculpturing indistinct; slightly inflated, usually broad-
est near level of hair 1-S; pecten teeth usually 7-11 (4-13); hair 1-S usually located
Q.25-0.33 (0.23-0.36) distance from base of siphon. Anal Segment: Saddle small,
brown, with sculpturing indistinct except apically; hair 1-X usually single (single,
double); gills sausage-shaped, dorsal and ventral subequal in development and usu-
ally 4.0-5.0 (3.8-5.5) length of saddle. |
Zavortink: A New North American Aedes 3
SYSTEMATICS. As now interpreted, the varipalpus complex of Aedes (Ochlero-
tatus) contains 4 species. Three of these, deserticola, monticola and varipalpus, are
relict species with relatively limited allopatric distributions in the interior portions
of the southwestern United States and in Baja California. The remaining species,
sierrensis, is very widely distributed in Pacific Coast drainage systems from British
Colombia to southern California and, in addition, is found along the desert margins
in western Nevada, northern Utah and southern California. While its distribution is
basically complementary to the other 3 species, it does occur with deserticola along
the desert edges in southern California.
All species in the varipalpus complex are very similar morphologically and are sep-
arated by relatively few characters in each stage. These diagnostic features are to be
found in the provisional keys to the complex given below.
BIONOMICS. Nothing is known of the habits of the adults of Ae. deserticola. The
larvae, like those of the other 3 species of the varipalpus complex, occur in treeholes.
On the northern side of the San Gabriel Mountains deserticola has been collected in
large rot holes in sycamores (Platanus racemosa) and a cottonwood (Populus fremon-
tii) growing in riverine situations. Ae. sierrensis occured in the same holes and Ortho-
podomyia signifera (Coquillett, 1896) was present at one of the localities. In Banner
Canyon deserticola has been recovered from large rot cavities in Engelmann oaks
(Quercus engelmannii). Ae. sierrensis was taken from the same holes and O. signifera
was present at the site. On the eastern side of the San Bernardino Mountains and in
the Little San Bernardino Mountains deserticola has been collected from small rot
holes in scrub oaks (Quercus turbinella) growing in pinyon-juniper woodland. Ae.
sierrensis and O. signifera were not found at these localities. The rot holes in the
limbs of these scrub oaks were remarkably small, most of them having an opening
less than 1 cm in diameter and holding less than 20 ml of water. In 2 instances larvae
of deserticola were removed from rot holes which had formed in a living limb be-
neath the base of a still-attached but dead and dried branch. In both cases the only
access to the cavity containing water was through the oval gallery of a wood-boring
beetle larva which had exited neat the base of the dead branch.
When larvae of sierrensis and deserticola are reared in the same container, the de-
velopment of deserticola is much slower than that of sierrensis. It is not known if
this is due to deserticola having an intrinsically slower rate of growth or due to its
being competitively inferior to sierrensis.
DISTRIBUTION. Ae. deserticola has been collected in the western portions of
both the Mojave and Colorado deserts, on the interior slopes of the Transverse and
Peninsular ranges and in the Little San Bernardino Mountains. During this study the
material cited below, all of which is in the UCLA collection, has been seen. Material
examined: 1321 specimens; 154 6, 128 9, 311 pupae, 728 larvae; 148 individual
rearings (138 larval, 10 incomplete).
CALIFORNIA. Los Angeles Co.: Littlerock (5 mi SW), type series, see above. Pearblossom (6.5
mi SE), 12 Mar 1969, T.J. Zavortink and D.W. Heinemann (UCLA 530), 1 Ipdé (530-12), 4 Ip?
(530-10,11,14,15), 1 6, 1 9, 2 P, 1 L. Pearblossom (8 mi SE), 4 Feb 1969, T.J. Zavortink (UCLA
518), 2 Ipd (518-21,25), 3 Ip? (518-28-30), 8 L. Riverside Co.: Joshua Tree (11-12 mi SSE), 17
Mar 1969, T.J. Zavortink and D.W. Heinemann (UCLA 539), 18 Ipd (539-10-27), 5 Ip? (539-28-
32), 17 6, 12 2, 33 P, 138 L; same data (UCLA 545), 4 Ipd (545-10,11,16,17), 3 Ip? (545-14,18,
19), 3 Ip (545-12,13,15),2 6, 4 9, 6 P, 4 L; same data (UCLA 546), 2 Ipd (546-10,12), 1 1p
(546-14), 3 lp (546-11,13,15). San Bernardino Co.: Yucca Valley (6.5 mi NW), 18 Mar 1969,T.J.
Zavortink and D.W. Heinemann (UCLA 544), 3 Ipd (544-10,11,13), 2 Ip? (544-14,15), 1 lp (544-
12); same data (UCLA 548), 1 Ipd (548-13), 4 lp? (548-10-12,14). Yucca Valley (9 mi NW), 18
Mar 1969, T J. Zavortink and D.W. Heinemann (UCLA 547), 1 L. San Diego Co.: Julian (34 mi
4
Contrib. Amer. Ent. Inst., vol. 5, no. 1, 1969
E), 12 Feb 1969, T.J. Zavortink and J.A. Bergland (UCLA 527), 13 Ipd ($27-23-25 ,27-32,34,37-
39), 15 lp? (527-22,26,33,35 ,36,40-49), 26 5,5 9, 30 P, 71 L; same data (UCLA 528), 2 Ipd (528-
20,21), 3 Ip? (528-22,24,25), 1 Ip (528-23).
2(1).
3(2).
2(1).
KEYS TO THE VARIPALPUS COMPLEX
ADULTS
Subspiracular area with several light bristles. . . . . . . . . varipalpus
Subspiracular area without bristles .
Postcoxal acea with scales 2 Sk ee a ee ek oN monticoh
Postvoxal area without scales «ake £0 a i oe
Base of hindtarsal segment 4 usually with a distinct white ring; metameron
usually with a small patch of scales; male proboscis with light scales re-
stricted to patch orring near middle .. . . . Slerrensis
Base of hindtarsal segment 4 usually dark or with only a few white scales
dorsally; metameron without scales; male proboscis with light scales in
long streak on at least ventral surface in basal half . . . . . deserticola
MALE GENITALIA
Basal tergomesal lobe of sidepiece with a dense patch of strongly developed,
apically curved setae; tergomesal margin of sidepiece with numerous short,
strong hairs; clasper not strongly curved apically, its spiniform relatively
short; connection between claspettes a horizontal but with a deep
median emargination .. . . . Slerrensis
Basal tergomesal lobe of sidepiece with a 1 single strongly differentiated seta;
tergomesal margin of sidepiece without hairs; clasper strongly curved api-
cally, its spiniform relatively long; connection between claspettes decliv-
Ouse res ee eer) Pee a en a
Basal tergomesal lobe of sidepiece with numerous long slender bristles in ad-
dition to | strongly differentiated seta . . . . varipalpus
Basal tergomesal lobe of sidepiece with 1-3 long slender bristles in addition
tO stronely dilterentiated Seta: 6s 2. A OW
Tergite [X usually with 4-6 (2-7) bristles on each lobe. . . . . deserticola
Tergite IX usually with 6-8 (5-10) bristles on each lobe . . . . monticola
PUPAE
Hair 1-IV relatively well developed, larger than 1-II] or 1-V. . . varipalpus
Hair 1-[V more weakly developed, usually not longer than 1-II] or 1-V . .2
Integument of dorsal portion of cephalothorax, metanotum and abdominal
segments 1-HI,[V or V of all but depauperate specimens well pigmented,
Tan dO DOW A eatin ee ee et ck SIRO RGIS
Zavortink: A New North American Aedes 5
Integument uniformly lightly pigmented, yellow. . ........ .3
3(2). Hair 5-VI usually oe in length to hair9-VIJ .. . . . . monticola
Hair 5-VI usually distinctly longer than hair9-VII . . . . . . deserticola
LARVAE
i: Body, especially dorsally and/or caudally, almost always pigmented, grey in
color; hair 1-IV,V usually shorter and frequently with more numerous
branches than hair 13 of the corresponding segment and usually shorter
than or subequal to hair 1-S; comb scales relatively numerous, usually 16-
24 (13-36), and arranged in 2 or 3 irregular rows or a small patch; pecten
teeth relatively numerous, usually 10-15 (7-17); siphon usually relatively
long, not noticeably inflated and not sharply reduced in diameter apically
ok ee a a ea ee a
Body usually unpigmented, white in color; hair 1-[V,V as above or subequal
in length to and with the same number of branches as hair 13 of the cor-
responding segment and longer than hair 1-S; comb scales few to numer-
ous (5-23) and arranged in 1-3 rows or a small patch; pecten teeth few to
numerous (4-18); siphon often relatively short, frequently noticeable in-
flated and/or sharply reduced in diameter apically . . ...... .2
2(1). Hairs 1,13-IV,V frequently subequal in length, usually double with 1 branch
much longer than the other and as long as the siphon; pecten teeth rela-
tively few, usually 7-11 (4-13); hair 1-X usually single, comb scales rela-
tively few, usually 7-12 (5-15), and usually arranged in a single regular or
irregular row; hairs 1,3-VIII frequently single . . . . . . . deserticola
Hair 1-[V,V shorter and with more numerous branches than hair 13 of the
corresponding segment, or if subequal in length to hair 13 and double,
then both hairs 1 and 13 shorter than siphon and usually without 1 branch
much elongate; pecten teeth relatively numerous, usually 10-17 (8-18);
hair 1-X usually double; comb scales few to numerous, usually 9-19 (7-
23), and arranged in 1 to 3 rows ora small patch; hairs 1,3-VIII branched
ee, 2
3(2). Hair 1-IV,V usually with the same number of branches as and frequently
subequal in length to hair 13 of the corresponding segment and longer
than hair 1-S; comb scales relatively numerous, usually 12-19 (7-23), and
usually arranged in 2 or 3 irregular rows or a small patch. . . monticola
Hair 1-IV,V usually much shorter than and with more numerous branches
than hair 13 of the corresponding segment and usually shorter than hair
1-S; comb scales relatively few, usually 9-14 (7-20), and usually arranged
Im.d OF 2 WregularToOWs 00k. 6 a ek
FIGURES
1. Aedes (Ochlerotatus) deserticola; male genitalia and pupa.
2. Aedes (Ochlerotatus) deserticola; larva.
o
oO H
Oo =s2
ee
9S
re S33
(cb) Osx
a BOR
(<b)
OO
AEDES
—Zy
a —_— be ZZ
deserticola
ihr ne is
alifor
Un ited States
rN EA
MOSQUITO STUDIES (Diptera, Culicidae)
XVII. TWO NEW SPECIES OF DEINOCERITES
FROM COSTA RICA’
By
Abdiel J. Adames” and Charles L. Hogue®
The new species, the first since the review of Belkin and Hogue (1959), are being
described here, detached from a general revision of the genus Deinocerites by
Adames, in order to make the names available for a forthcoming publication by
Hogue and Donald B. Bright, California State College, Fullerton, on the biologies of
land crabs and their burrow associates in Costa Rica. Most of the material of the new
species was collected as part of a worldwide study of tropical land crabs and their
burrow associates (LCBA) being conducted by Hogue and Bright with the support
of 2 grants from the American Philosophical Society. Additional specimens were ob-
tained through the project ‘“Mosquitoes of Middle America” and also from the pro-
ject “A Study of the Mammalian Ectoparasites and their Hosts in Costa Rica’’, the
latter supported by the U.S. Army Medical Research and Development Command
(DA-MD-49-193-62-G54,G94).
The method of presentation, terminology and abbreviations used in the descrip-
tions follow Belkin (1962), whom we thank for advice. In the figures of immature
stages, the hair branching shown is based on the modal values for 8 specimens of
nicoyae and 10 specimens of costaricensis.
Deinocerites nicoyae Adames & Hogue, n.sp.
Figs. 1-3
TYPES: Holotype 6 with associated larval and pupal skins and genitalia slide (CR 254-21),
Estero El Mero, Boca del Rio Barranca, Puntarenas Province, Costa Rica, 11 Feb 1969, C.L. Hogue
and DB. Bright [USNM] . Allotype ? with associated pupal skin (254-100), same data as holotype
[USNM] . Paratypes: 5 \pd (CR 254-23,25 ,32,34,49), 2 pd (254-101,104), 2 IP (254-18,52), 157
L, 1 1(254), same data as holotype [BM,LACM,UCLA,USNM].
‘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.
* Entomology Section, Los Angeles County Museum of Natural History, Los Angeles, California
90007.
10 Contrib. Amer. Ent. Inst., vol. 5, no. 1, 1969
FEMALE (fig. 1). Wing: 2.79 mm. Proboscis: 1.69 mm. Forefemur: 1.61 mm.
Abdomen: about 2.83 mm. Small species. Head: Integument uniformly dark; narrow
decumbent scales of vertex dark brown; erect scales brown to dark brown, lateral
patch of scales whitish. Clypeus dark. Labium and palpus dark scaled. Antenna dark;
flagellar segment 1 with scales, subequal to combined length of segments 2, 3 and
two-thirds of 4, other segments not markedly elongate; proboscis extending to 8th
flagellar segment; torus dark. Thorax: Mesonotal integument dark, strongly contrast-
ing with very distinctly lighter portions of the pleural integument. Postnotum usual-
ly with a few hairs. Apn with light brown integument anteriorly, remainder light;
upper ppn with dark scales and bristles, middle and lower areas light and with | to 3
small hairs; ppl with brown integument and several bristles; psp and ssp with dark
brown integument, bare; stp with dark brown integument, a patch of translucent
scales and a row of bristles along dorsal and caudal margins; paratergite dark and
bare; pra with lighter integument and several bristles; mep integument light and with-
out scales, umep with several bristles, Imep with 1 very strong bristle; 4 or 5 short
hairs below base of haltere; meron, metameron and metapleuron light. Legs: Anter-
ior surface of forecoxa largely covered by bristles and translucent scales; midcoxa
with lateral surface barely darker than adjacent portions of pleuron and with several
scales and 2 rows of bristles, the posterior bristles stronger; hindcoxa with lateral
surface similar in color to adjacent portion of pleuron and with a few scales and
bristles, posterior surface with a row of bristles; femora, tibiae and tarsi entirely
dark scaled. Forefemur without spiniforms but with 4 to 6 bristles on anteroven-
tral distal part and with a row of light bristles on posterodorsal side. Wing: Vein and
fringe scales dark. Haltere: Integument of stem light, knob dark scaled. Abdomen:
Tergites uniformly dark scaled; sternites II-VII about the same color as tergites.
FEMALE GENITALIA (fig. 1). Sternite VIII with a few scales on sclerotized api-
cal part. Tergite IX with 1 or 2 small setae on each side. Cercus without spatulate
apical or subapical specialized setae. Postgenital plate with 1 apical specialized seta.
MALE (fig. 1). Wing: 2.91 mm. Proboscis: 1.90 mm. Forefemur: 1.90 mm. Abdo-
men (not including genitalia): about 1.67 mm. Similar to female in general colora-
tion. Flagellar segments 1-3 with dark scales; segments 1-6 markedly elongated; seg-
ment 13 expanded subapically; segment 1 shorter than 2 and 3 combined; pro-
boscis not extending beyond middle of segment 5. Claws of foreleg and midleg
sek anterior claws with a heavy submedian tooth, posterior claws without
ooth.
MALE GENITALIA (fig. 2). Segment [X: Tergite lobe cylindrical, broad and ang-
| led laterad at base; distal part reaching level of subapical lobe (median mesal lobe of
Belkin and Hogue, 1959), directed inwards distally by a distinct deep mesal curva-
ture or constriction, expanded apically. Sidepiece: Without scales. Subapical lobe
with thumb; ventromesal surface with 3 well-differentiated setae, the most anterior,
in dorsal aspect, bristle-like and attenuated apically, the 2 posterior spiniforms rather
heavy and without apical attenuation. Apicosternal lobe long, with differentiated
apical seta. Phallosome: Dorsal parameres widely separated in tergal aspect, caudo-
ventral margin expanded laterad as a broad hemispherical ledge, without sclerotized
dorsal bridge; apical spine short, heavy and strongly curved dorsad, visible in toto
only in lateral aspect; ventral teeth large, heavy, progressively shorter toward apex
and not arising from a distinct lobe. Aedeagus cylindrical, broad mesally then con-
hea apex poorly sclerotized but well defined by a subapical necklike constric-
ion.
PUPA (fig. 2). Abdomen: 2 3.56 mm; 6 3.34 mm. Trumpet: 0.51 mm. Paddle:
Adames & Hogue: Two New Deinocerites 11
0.71 mm. Chaetotaxy as figured. In general similar to other members of the genus;
differing in the following diagnostic characters. Cephalothorax: Hair 5-C double,
strongly developed, longer than distance from its alveolus to base of trumpet, about
1.3 length of trumpet; hair 8-C long, single; hair 9-C short, about one-third to nearly
half length of 8-C. Trumpet: Index about 4.5-4.6; pinna short. Metanotum: Hair 10-
C short, smaller than 11-C, usually single. Abdomen: Tergal area between hairs 1-I
without contrasting pigmentation. Hair 1-II not reaching apex of tergite III, triple,
the middle branch longest; hair 5-IV,V barely exceeding the base of succeeding seg-
ment; hair 1-VII subequal to tergite VIII. Paddle: Width about two-thirds of length;
paddle hair (1-P) usually as long as paddle.
FOURTH INSTAR LARVA (fig. 3). Head: 1.05 mm. Siphon: 1.14 mm, index
about 4.7-5.3. Chaetotaxy as figured. Similar to other members of genus; with the
following diagnostic features. Head: Mental plate slightly wider than long, with
sharply pointed marginal spicules. Hair 2-C inconspicuous, about 0.2 length of 1-C,
either slightly mesad, in line with or laterad of level of 1-C; 3-C not detectable; hair
5-C longer than antenna, usually 4 branched (2-5); 6-C longer than 5-C, single, barb-
ed. Antenna: Length about 0.33 of head, shaft with a few minute spicules on proxi-
mal part. Thorax: Hair 3-P always double; 8-P double; 9-P single; 4-M usually triple
(2-4); 6-T short, usually double. Abdomen: Hair 3-I longer than 4-I; 6-I-V strong, al-
ways double; 6-VI strong, double, branches unequal; 1-[V moderately long, exceed-
ing base of segment V, usually single; 1-VII short, smaller than 3-VII and not reach-
ing base of segment VIII; 1-VIII usually 6 branched (5-7). Siphon: As figured; hair
1-S relatively short, triple, branches unequal; pecten with 4 to 6 teeth on each side.
Anal Segment: Hair 1-X short, double to 5 branched; hair 2-X usually with 5 to 7
branches. Ventral brush (4-X) with 6 pairs of hairs. Gill short, slightly emarginate on —
apex.
SYSTEMATICS. On the basis of several features of the adults, nicoyae falls into
the Dyari Group as defined by Belkin and Hogue (1959). The female of this species
is apparently indistinguishable from that of dyari Belkin & Hogue, 1959, the only
other known species of this group. The male of nicoyae can be distinguished from
dyari by the IX tergite lobe reaching the level of the subapical lobe while in the lat-
ter itis short and does not reach the level of the lobe. The immature stages of nicoy-
ae can be readily separated from dyari (at least from Colombian populations) by the
following features: (1) in the pupa, hair 5-C double instead of single; hair 1-II triple
instead of single; (2) in the larva, ventral brush (4-X) with 6 pairs of hairs instead of
7 pairs; hair 6-III double instead of 3 or 4 branched.
BIONOMICS. D. nicoyae has been collected only in the burrows of the Wide Red
Land Crab, Ucides occidentalis (Ortmann), family Ocypodidae. The immature stages
were found on | occasion only, when the water level in these burrows was consider-
ably depressed. Occasionally associated with nicoyae was D. pseudes Dyar & Knab,
1909 which more frequently utilizes the burrows of Cardisoma crassum Smith (Gec-
arcinidae).
DISTRIBUTION. Known to date only from the type locality in Costa Rica and
about 3 miles west of same. Material examined: 418 specimens; 151 6,92 9, 11 pu-
pae, 167 larvae, 11 individual rearings (3 pupal, 6 larval, 2 incomplete).
COSTA RICA. Puntarenas: Boca del Rio Barranca, Estero El Mero, 6 July 1967, C.L. Hogue
(CR 238), 2 6, 10 ?, 10 July 1967, C.L. Hogue (CR 240), 58 6, 24 9 [UCLA]; 10 July 1967:
C.L. Hogue (LCBA-188), 69 6, 53 @ [LACM]; 11 Feb 1969, C.L. Hogue and D.B. Bright (CR
254), 6 Ips (254-21,23,25,32,34,49), 2 pd (254-101,104), 1 p? (254-100), 2 IP (254-18,52), 157
L, 1 1 (254) [Type series, USNM,BM,LACM,UCLA]. La Angostura, 7 Nov 1962, JN. Belkin,
C.L. Hogue and W.A. Powder (CR 3); 6 6, 3 9, 17 Nov 1962, J.N. Belkin, CL. Hogue and W.A.
Powder (CR 27), 8 6, 1 9 [UCLA].
12 Contrib. Amer. Ent. Inst., vol. 5, no. 1, 1969
Deinocerites costaricensis Adames & Hogue, n.sp.
Figs. 4-6
TYPES: Holotype ? with associated larval and pupal skins (CR 28-213), 1 km north of Boca
del Rio Barranca, Hacienda Bonilla, Puntarenas Province, Costa Rica, 17 Nov 1962, C.L. Hogue
and W.A. Powder [USNM]. Allotype 6 with associated larval and pupal skins and genitalia slide
(CR 28-210), same data as holotype [USNM] . Paratypes: 3 lp? (CR 28-214,215,218), 3 pd (28-
201,203,205), 1 p? (28-204), 1 IP (28-216), 5 6, 3 ?, 113 L, 3 P, 5 p (CR 28), same data
as holotype [BM,LACM,UCLA,USNM]}.
Deinocerites species A of Belkin and Hogue (1959:438); Hogue and Wirth (1968:6).
FEMALE (fig. 4). Wing 3.98 mm. Proboscis: 2.37 mm. Forefemur: 2.24 mm.
Abdomen: about 4.40 mm. Medium-sized species. Head: Integument markedly dark;
narrow decumbent scales on vertex creamy; erect scales yellowish; lateral patch of
scales whitish. Clypeus dark. Antenna with numerous scales on flagellar segment |
and a few on segment 2, segments 1 to 4 greatly elongated, segment 1 about equal
to combined length of segments 2 and one-third of 3; proboscis not extending be-
yond middle of segment 5; torus dark, with a few short hairs on mesal surface. Thor-
ax: Mesonotal integument dark brown. Postnotum without hairs. Apn with tan col-
ored integument, anteriorly with a row of strong bristles and posteriorly with a
transverse row of less developed bristles; ppn tan colored, upper ppn with scales and
bristles, middle and lower areas with bristles; pp! with tan colored integument and
numerous bristles; psp and ssp with brownish integument, bare; stp with tan colored
integument, completely covered by translucent scales, a row of bristles arranged
along dorsal and caudal margins; paratergite brownish and bare; pra with several
bristles; mep with tan colored integument and covered by a patch of translucent
scales, umep with several bristles, Jmep without a long bristle; a few short hairs at
the base of haltere; meron, metameron and metapleuron with tan colored integu-
ment, metameron with several small hairs. Legs: Anterior surface of forecoxa largely
covered by bristles and scales; midcoxa of the same color as adjacent portions of
pleuron, outer surface extensively covered with translucent scales and 2 rows of
bristles, the posterior bristles stronger; hindcoxa with integument similar in color to
adjacent portions of pleuron, anterolateral surface with translucent scales, lower lat-
eral surface with bristles, upper lateroposterior surface with scales, posterior surface
with numerous bristles; femora, tibiae and tarsi dark scaled; forefemur with an an-
teroventral and posterodorsal row of bristles not modified into spiniforms; hind-
tibia with a row of spiniforms on the apical two-thirds of dorsal surface. Wing: Vein.
and fringe dark scaled. Haltere: Stem light, dorsoapical part with several dark scales;
knob dark scaled. Abdomen: Tergites dark scaled; sternites II-VII lighter than tergites.
FEMALE GENITALIA (fig. 4). Sternite VIII without scales on sclerotized apical
part. Tergite IX with a pair of small setae on each side. Cercus with 6 apical and sub-
apical specialized setae with recurved and twisted apex. Postgenital plate with 1 api-
cal specialized seta.
MALE (fig. 4). Wing: 3.13 mm. Proboscis: 2.71 mm. Forefemur: 2.37 mm. Abdo-
men (not including genitalia): about 2.79 mm. Similar to female in general colora-
tion. Flagellar segments 1-4 with scales, segments 1-6 markedly elongated; segment
1 3 slightly expanded subapically; segment 1 shorter than 2 and 3 combined; probos-
cis not extending beyond the basal part of segment 4. Anterior claw of foreleg with
a very long, slender, subbasal tooth, posterior with a minute subbasal projection; an-
Adames & Hogue: Two New Deinocerites 13
terior claw of middle leg with a minute subbasal tooth, posterior claw simple.
MALE GENITALIA (fig. 5). Segment IX: Tergite lobe cylindrical, angled laterad
from base, broad at base and cone-shaped distally, apical part not strongly attenu-
ated and not reaching level of subapical lobe. Sidepiece: Without scales. Subapical
lobe with rather small thumb, seta c spiniform, with an apical attenuation; ventro-
mesal surface with 3 distinct bristlelike setae, attenuated apically. Apicosternal lobe
with differentiated apical seta. Phallosome: Dorsal parameres widely separated in ter-
gal aspect but with a slight indication of an incomplete dorsal bridge; apical spine
long and slender. Aedeagus cylindrical, more or less uniform in width but with a sub-
median constriction and an apical expansion on sclerotized part.
PUPA (fig. 5). Abdomen: ? 4.07 mm, 6 3.62 mm. Trumpet: 0.76 mm. Paddle:
0.84 mm. Chaetotaxy as figured. Similar to other members of genus, with the fol-
lowing diagnostic characters. Cephalothorax: Hair 5-C single, moderately long, equal
or subequal to distance from its alveolus to base of trumpet, subequal in size to
length of trumpet; 4-C single, more than half length of 5-C; 8-C usually triple, sub-
equal to length of trumpet. 7Jrumpet: Index about 5.3-7.6; pinna small. Metanotum:
Hair 10-C well developed, longer than 11-C, single. Abdomen: Tergal area between
hairs 1-I without contrasting pigmentation. Hair 1-II longer than tergite III, always
exceeding the base of tergite IV, double or triple; 5-II mesad to 2-II, hair 5-III-V
long, extending as far as middle of second tergite following; hair 1-VII subequal to
tergite VIII. Paddle: Width more than two-thirds of length; paddle hair (1-P) longer
than paddle.
FOURTH INSTAR LARVA (fig. 6). Head: 1.22 mm. Siphon: 1.43 mm, index
about 3.8-5.6. Chaetotaxy as figured. In general similar to other members of genus,
with the following diagnostic characters. Head: Mental plate almost as wide as long
and with blunt apical and subapical marginal spicules. Hair 2-C inconspicuous, about
0.25 of length of 1-C, always mesad of level of 1-C; 3-C not detectable; hair 5-C long-
er than antenna, always double; 6-C longer than 5-C, always single and simple. An-
tenna: Length about 0.33 of head; shaft with numerous minute spicules in proximal
part. Thorax: Hair 3-P always single; 8-P usually single (1-2); 9-P usually triple (2-3),
4-M usually single (1-2); 6-M moderately long, single. Abdomen: Hair 3-I longer than
4-I; 6-I-V strong, double; dorsal sensillum of segment V mesad to hair 4-V; 6-VI
strong, single; 1-VII long, longer than 3-VII and exceeding base of siphon; 1-VIII
usually 3 branched (3-4); 3-VIII usually 4 branched (4-5). Siphon: As figured; hair 1-
S moderately long, reaching hair la-S, double, branches equal; pecten with 4 to 6
teeth on each side. Anal Segment: Hair 1-X short, usually double (1-3); hair 2-X usu-
ally 9 branched (6-10). Ventral brush (4-X) with 7 pairs of hairs. Gill long, subequal
to dorsal saddle length, slightly emarginate on ventral margin.
SYSTEMATICS. D. costaricensis is species A of Belkin and Hogue (1959), which
was placed by them in the Epitedeus Group. Its closest relative is epitedeus (Knab,
1907), which is now known to have a wide distribution along the Caribbean coasts
from the Gulf of Honduras to Colombia. The 2 species can be differentiated more
easily in the adults than in the immature stages. In costaricensis the forefemur has an
anteroventral row of bristles which in epitedeus are replaced by spiniforms. The male
of costaricensis can be differentiated from epitedeus by the IX tergite lobe being
short, cone-shaped and not reaching the level of subapical lobe while in the latter it
is long and slender distally and extends beyond the subapical lobe. No diagnostic
features have been found so far for the pupa. In the larva, costaricensis can be differ-
entiated from epitedeus by the following features: (1) hair 7-II is usually 5 branched
(3-7) instead of usually double (1-4), (2) the mental plate generally with the term-
inal spicules blunt instead of sharply pointed.
14 Contrib. Amer. Ent. Inst., vol. 5, no. 1, 1969
BIONOMICS. All records of costaricensis are from the burrows of the Mouthless
Crab, Cardisoma crassum Smith (Gecarcinidae). This species appears to be rare com-
pared to pseudes which also utilizes these burrows.
DISTRIBUTION. This species is known only from the Pacific coast of Costa Rica;
however it has not been found in collections of Deinocerites from the outer coast of
the Nicoya Peninsula. It may extend farther south along the Pacific coast of Panama.
Material examined: 689 specimens; 31 6, 41 ?, 89 pupae, 528 larvae, 38 individual
rearings (5 pupal, 23 larval, 10 incomplete).
COSTA RICA. Puntarenas: Boca del Rio Barranca, 31 July 1962, F.S. Truxal, 2 d, 3 9; 27 June
1967, C.L. Hogue and D.B. Bright (LCBA 111), 3 L; 8 July 1967, C.L. Hogue and D.B. Bright
(LCBA 155,156,158-159), 6 L; 11 July 1967, C.L. Hogue and DB. Bright (LCBA 174), 1 9
[UCLA]. Hacienda Bonilla, 1 km north of Boca del Rio Barranca, 17 Nov 1962, C.L. Hogue and
W.A. Powder (CR 28), 3 pd (28-201,203,205), 1 p? (28-204), 1 lpd (28-210), 4 Ip? (28-213,214,
215,218), 1 IP (28-216),5 6,3 9,111 L,3 P (28-2) [type series, USNM,BM,LACM,UCLA] ; 20 June
1963, C.L. Hogue (CR 106,107), 1 ¢ (106), 16 L (107); 21 June 1963, C.L. Hogue (CR 118), 2 9.
Boca del Rio Baru, 3.5 km northwest, 20 Nov 1962, C.L. Hogue and W.A. Powder (CR 34,43), 2
Ips (34-109,114), 5 Ip? (34-105,106,110,112,115), 1 p? (34-101), 6 IP (34-102,103,104,108,
111,113), 6 6,9 2,210 L, 51,8 P, 16 p (34), 1 6 (43) [UCLA] . Rincon, Peninsula de Osa, 26 June
1963,C.L. Hogue(CR 122-124), 36 L, 1 P (122), 2 Ipd (123-101,109), 4 Ip? (123-102,105-107), 1
IP (123-104), 4 6,2 2,9 P, 5 p, 47 L, 41(123), 1 6 (124); 29 June 1963, C.L. Hogue (CR 130,135-
136), 1 1 (130, mangrove treehole), 2 Ipd (135-105,107), 1 IP (135-106), 4 L (135), 2 lpd (136-
101,103), 1 Ip? (136-102), 1 IP (136-104); 2 July 1963, C.L.-Hogue (CR 142), 5 6,29,54L,1
1,4 p (142) [UCLA]. Tarcoles, 1 9 [USNM; Belkin and Hogue, 1959:438, as species A].
REFERENCES CITED
Belkin, John N. |
1962. The mosquitoes of the South Pacific (Diptera, Culicidae). v. 1. Berkeley,
University of Calif. Press. 608 p.
Belkin, John N. and C.L. Hogue
1959. A review of the crabhole mosquitoes of the genus Deinocerites (Diptera,
Culicidae). Calif. Univ. Publ. Entomol. 14:411-458.
Hogue, Charles L. and W.W. Wirth
1968. A new Central American sand fly breeding in crab holes (Diptera, Cerato-
pogonidae). Los Angeles County Mus., Contrib. Sci. 152.7 p.
FIGURES
1. Deinocerites nicoyae; male and female heads, female genitalia, fore and mid- ©
claws of male, forefemur of female
2. Deinocerites nicoyae; male genitalia and pupa
3. Deinocerites nicoyae; larva
4. Deinocerites costaricensis; male and female heads, female genitalia, fore and
midclaws of male, forefemur of female
5. Deinocerites costaricensis; male genitalia and pupa
6. Deinocerites costaricensis: larva
DEINOCERITES
e C rf 5 KEE Zz
——— y TP irs ey
= SQ EEE
—_. — s Whe ee pahee
—
——— eee
post genital plate
:
spermatheca
anterior posterior
Fig. |
FORE
J 0.2
nicoyae
LCBA 188
anterior posterior
N
iN MID
ee ———caten errno aan earner
left anterior
Pare a ST eT are
right posterior
rer
dorsal paramere
me res
nicoyae
CR 254
DEINOCERITES
— = es
a)
=
tip x
nicoyae
CR 254
DEINOCERITES
anterior posterior
See
>
FORE
=
Hin
a i
Ke anterior posterior
Lg
AN
A costaricensis MID
oe BR CR 34
i
i.
HAIN
| oe
aed e) left anterior
in |
ae ae oe ee a
right posterior
pe Se ee Oger ere ee
costaricensis
”
Li]
=
or
Li
S
O
=
i
a
DEINOCERITES
eee
3
i
1\
1
fl
tS y | /
> 2
i}
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11
costaricensis
CR 34
MOSQUITO STUDIES (Diptera, Culicidae)
XVIII. THE SUBGENUS MICRAEDES OF CULEX’
By
O.G.W. Berlin?
CONTENTS
INTRODUCTION _. iri ee a ee ee a a ig
GENERAL CONSIDERATIONS OS UE ae ae ee ee
TexOnomiet Maractere es ee A ee
DORITONE OR Lae Ve re a eee a ee
DistiDntion oF ao ee ee a re
Dy lematiCs Src a SG ee ie ei re Ee eg ee
Affinities. . . ee Ce ee a a ae ee
TAXONOMIC TREATMENT . Pee ee ee ye ee ee ee
Subgenus Micraedes. . . Le ee er an ee ee
Keys to Groups and Species Prt, Oa ee Mae ee ee
Bisulcatus Group . POP ee err ee, ay
1. Culex LW) bisiiledtils ( Coquillett) SEAS ar gee en tee er tee oe
2. Culex (MM. Jantilwnmagnoriin Dyar (6 OPO Oe aa
See A eae a Pe Oe Wa ee es De 99
SchickiGroup . . A en er i ee ee re ee
4. Culex (M. ) schicki, n. sp. ge SR CGE TONE PTET tr ce ae
en (EF nde. WS BO ON Ss agg
Frethyzonfer Group, SERN Deg GOA! SEE ORC ern as
6. Culex (M. ) erethyzonfer Galindo & Blanton ae i viras Gana ce |.
Ber Ce Cee re rs Re Pe ee
ee ee er ee ee
PIONS PORMCTEINT LIC BANE ee teen ae SE I UP eae
INTRODUCTION
Micraedes was proposed as a distinct genus by Coquillett (1906a:185) with the
new species bisulcatus as type and with the following definition: ‘““Near Aedes, but
the palpi much longer in both sexes. Palpi noticeably longer than the head, about
‘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. Permanent
address: Department of Zoology, American College, Madurai, Madras State, India.
22 Contrib. Amer. Ent. Inst., vol. 5, no. 1, 1969
one-third as long as the proboscis, apparently two-jointed, the apical joint slightly
longer than the basal. Hairs of male antennae much longer and more numerous than
in the female. Clypeus bare. Occiput covered with narrow, curved scales and with
many upright forked ones, the lateral portions covered with very broad, appressed
scales. Scutellum bearing very narrow scales and six bristles on the median lobe.
Scales of legs appressed. Venation normal, hind crossvein over twice its length be-
fore the small, the lateral scales of the veins very narrow.’ Howard, Dyar and Knab
(1915:216) synonymized Micraedes with Culex but Dyar (1918:102) elevated it to
the subgeneric rank within Culex. In 1928 Dyar synonymized bisulcatus with ameri-
canus (Neveu-Lemaire, 1902) which was placed in the subgenus Melanoconion, and
as a consequence, Micraedes became a synonym of Melanoconion. Subsequently Ed-
wards (1932:218) restored Micraedes to subgeneric rank, Lane (1953:387) synony-
mized it with the subgenus Tinolestes and Stone, Knight and Starcke (1959:281)
with the subgenus Aedinus. :
In the present study, Micraedes is again treated as a distinct subgenus as has been
done recently by Foote (1954:4) and Belkin (1968:11). In addition to bisulcatus,
which was first correctly removed from synonymy with americanus by Floch and
Abonnenc (1945:37), it now contains 5 other species which appear to be related to
bisulcatus on the basis of significant correlated features of the adults and immature
stages. These species fall into 3 well defined groups. |
I wish to express my gratitude to John N. Belkin for suggesting this study and for
examining the types in the U.S. National Museum and the British Museum. I am in-
debted to Alan Stone of the U.S. National Museum for the loan of material and
to Kenneth L. Knight for the loan of C. antillummagnorum from Cuba and Puerto
Rico. Finally, I wish to thank Sandra J. Heinemann, William A. Powder and Abdiel
J. Adames for valuable assistance, Nancy Martsch for the preparation of the final
plates of male genitalia and Sheila Bernstein and Caryle Abrams for the preparation
of the copy for reproduction.
This study is based primarily on material collected and accumulated at the Univer-
sity of California, Los Angeles, for the project “Mosquitoes of Middle America”’
(Belkin, Schick et al, 1965) and also on material from the U.S. National Museum.
Nearly 2000 specimens (189 6, 196 2, 367 pupae, 1226 larvae) including 304 in-
dividual rearings (65 pupal, 147 larval, 92 incomplete) were examined. The method
of presentation and the terminology and abbreviations used in the descriptions in
general follow Belkin (1962). In the illustrations, all available stages of every species
are figured, the larvae and pupae in full and the adults in pertinent details only. In
the distribution lists, the arrangement of countries, major political subdivisions and
localities is alphabetical throughout. The abbreviations used in the synoptic catalog |
(Stone, Knight and Starcke, 1959) for institutions in which types are deposited are
followed here to indicate the location of the material studied. To these is added the
Department of Zoology, University of California, Los Angeles [UCLA] where the
bulk of the material will remain. The abbreviations used for the references follow
the “‘American Standard for Periodical Title Abbreviations” (Amer. Stand. Ass.,
1964). The types of all new species described here will be deposited in the U.S.
National Museum.
GENERAL CONSIDERATIONS
TAXONOMIC CHARACTERS. Adults of Micraedes, like those of most other-sub-
genera of Culex, are very similar in external features and therefore I have not been
Berlin: Subgenus Micraedes of Culex 23
able to find many reliable characters for the separation of groups. At the specific
level the problem is acute, as there is no way of separating the different members of
a given group except geographically. The few but significant characters that I found
useful in the separation of groups are: (1) length and/or segmentation of the palpus
of males and females, (2) coloration of scales on the upper margin of ppn, and (3)
presence or absence of a bristle on lower mep.
The male genitalia show some significant and reliable characters at group and spec-
ific levels as follows: (1) number and length of bristles on the ninth tergite lobe, (2)
position and details of the subapical lobe of the sidepiece, (3) details of the clasper,
(4) presence or absence of foliform setae on the subapical lobe, and (5) number of
apical spines on the paraproct.
The immature stages, on the other hand, show an array of characters at the sub-
generic level separating Micraedes from other subgenera of Culex and significant fea-
tures to separate the various groups and the individual species. In the pupae the char-
acters used at the group level are: (1) length of cephalothoracic hair 5-C, (2) position
of abdominal hair 2-VII, (3) length of abdominal hair 9-VIII in relation to segment
VIII, and (4) presence or absence of spicules on the paddle margin. The fourth instar
larvae show many taxonomic characters at the group and specific levels. Among the
important characters used in the separation of groups are: (1) position of head hair
4-C, (2) development of basal maxillary hair (bmh), (3) development of hairs on the
thorax and abdomen (stellate or multiple), (4) position of abdominal hair 1 in rela-
tion to hair 2 on segments I-VII, (5) presence or absence of spines on the saddle mar-
gin, and (6) branching of saddle hair 1-X.
BIONOMICS. All members of the subgenus are leaf axil breeders, primarily in epi-
phytic and terrestrial bromeliads and rarely in aroids. They are usually associated
with species of Anopheles (Kerteszia), Aedes (Howardina), Culex (Microculex), Tox-
orhynchites (Lynchiella) and Wyeomyia. The records of antillummagnorum from
artificial containers in Cuba (Pazos) and Puerto Rico (Tower) mentioned in Howard,
Dyar and Knab (1915:306) have not been confirmed and are probably erroneous.
Very little is known of the bionomics of the adults of Micraedes. The only records
of blooded females are for antillummagnorum in Cuba and bisulcatus in Guadeloupe.
Both sexes have been taken resting during the day on tree trunks (antillummagnor-
um) or on moist soil in limestone caves (arawak), suggesting that Micraedes species
are crespuscular or nocturnal in activity. The record of antillummagnorum flying
and biting during the day (Pazos in Howard, Dyar and Knab, 1915:306) is probably
erroneous.
DISTRIBUTION (fig. 1). Micraedes is confined to the American Mediterranean
region in the sense of Belkin (1962:562). As far as known, the 3 groups of the
subgenus have an allopatric distribution. The Bisulcatus Group is restricted to the is-
lands of the Caribbean without any encroachment on the mainland; the Schicki
Group has been found only in the Sierra Madre del Sur in southern Mexico; and the
Erethyzonfer Group is known only from the mountain chains of Central America,
from Guatemala southward to Panama.
SYSTEMATICS. The following combination of characters of the adults and the
immatures will distinguish Micraedes from other subgenera of Culex. In the adults,
the subgenus is characterized by the presence of (1) only narrow decumbent scales
on the vertex, (2) acrostichal bristles on the mesonotum, (3) generally yellowish
pleural integument, (4) dark tarsi, and (5) basolateral pale patches on the abdominal
tergites. In the male genitalia, Micraedes is distinguished by (1) the shape of the
ninth tergite lobe, (2) the development and the position of the subapical lobe of the
sidepiece, and (3) the shape of the lateral plate of the phallosome. In the pupal stage,
24 Contrib. Amer. Ent. Inst., vol. 5, no. 1, 1969
species of Micraedes are diagnosed by (1) hair 5-C strongly developed, at least sub-
equal to length from its base to that of trumpet, (2) hair 2-VII always within 0.8 of
its tergite, (3) hair 9-VIII long, triple, either subequal to or distinctly longer than ter-
gite VIII, (4) absence of hair 1-[X, (5) presence of long, hairlike marginal spicules on
the paddle (absent in Bisulcatus Group), and (6) presence of only 1 terminal paddle
hair. Many diagnostic features in the larvae readily separate members of Micraedes
from other subgenera. Among the most important are: (1) absence of head hair 3-C,
(2) head hairs 5-7-C always multiple, long, subequal in length and at least 4 branch-
ed, (3) hair 1-A about 0.5-0.6 from antennal base, (4) mental plate with 7 well-devel-
oped lateral teeth, (5) many distinct stellate hairs on the thorax and the abdomen
(multiple in Schicki Group), (6) siphon with 4 pairs of well developed subventral
and 2 pairs of subdorsal hairs, (7) apical hook (2-S) of siphon slightly curved at tip
and with a subbasal branch, (8) caudolateral border of saddle with long marginal
spines, (9) saddle hair 3-X long, at least double, branches subequal, and (10) ventral
brush with only 5 pairs of 2-4 branched hairs.
The 6 species recognized in the subgenus Micraedes fallinto 3 well-defined groups:
(1) Bisulcatus Group with 3 species (bisulcatus, antillummagnorum and arawak), (2)
Schicki Group with 2 species (schicki and sandrae), and (3) the monotypic Erethy-
zonfer Group.
AFFINITIES. Micraedes is undoubtedly a member of the Melanoconion complex
of subgenera which includes also Microculex and a number of distinct phylads with a
short palpus in the male currently listed under Eubonnea and Aedinus (Stone,
Knight and Starcke, 1959:280-282), that Belkin (1968:11-12) suggests segregating
into Aedinus (~Eubonnea), Tinolestes and Anoedioporpa, and Micraedes. All of
these taxa have the aedeagus of the male genitalia essentially similar in structure but
each group is strongly differentiated in the immature stages and in ecology (breeding
sites). The short male palpus is not a good subgeneric character in the New World
Culex for as pointed out by Belkin (1968:11) ‘“‘this character occurs independently
in several unrelated phylads” and some species with a short palpus are unquestion-
ably related to species with a long palpus. This is especially true of Micraedes in
which the male palpus ranges in length from 0.35 to as long as the proboscis in the
different species.
TAXONOMIC TREATMENT
Subgenus MICRAEDES Coquillett
1906. Micraedes Coquillett, 1906a:185. TYPE SPECIES. Micraedes bisulcatus Coquillett, 1906,
Guadeloupe; original designation.
Culex (Micraedes) of Dyar (1918:102; 1922:95; 1923:187); Bonne and Bonne-Wepster (1925:
274), Edwards (1932:218-219); Lane (1939:76; 1949:255); Foote (1954:4); Belkin (1968:11).
Culex (Aedinus) in part of Stone, Knight and Starcke (1959:281); Belkin (1962:179); Stone
(1961:47; 1963:135; 1967:218); Belkin, Schick and Heinemann (1965:13); Forattini (1965:
ST 32 ,34,395165,193).
Culex (Tinolestes) in part of Lane (1953:387); Duret and Damasceno (1955:395).
Culex (Melanoconion) in part of Dyar (1928:343).
Culex in part of Howard, Dyar and Knab (1915:216).
Micraedes of Coquillett (1906b:24; 1910:569); Theobald (1910:486).
FEMALES (fig. 2). Small to medium sized inornate species; integument dark brown
Berlin: Subgenus Micraedes of Culex 25
to blackish except for pleuron which is usually yellowish; legs unbanded. Head: Eyes
not distinctly separated between antennae. Decumbent scales narrow and linear dor-
sally, broader ones restricted to sides and venter. Erect scales on occiput moderately
long, forked apically, extending to sides of vertex. Orbital and interorbital bristles
strong; upper orbitals usually 5(5-6) pairs, heavier, longer and more widely spaced
than lower. Clypeus bare, light to dark brown. Proboscis distinctly longer than or
subequal to forefemur; entirely dark scaled, with a few basal bristles. Palpus short to
moderately long, about 0.2-0.38 of proboscis; 4-segmented, segments 1 and 2 anky-
losed, without scales; segment 4 longer than 3, both dark scaled. Antenna subequal
to proboscis; torus light to dark brown, with a few short, dark brown setae mesally;
flagellar segments 2-13 with 6 moderately long bristles in basal whorls. Thorax: Inte-
gument moderate to deep brown. Mesonotum with short, narrow, curved, auburn to
dark scales except along a pair of inner dorsocentral “‘bare”’ lines extending from an-
terior margin to 0.75 of its length; light scales when present, restricted to anterior
promontory and humeral areas. Acrostichal bristles distinct, usually extending to
prescutellar area; dorsocentrals, prescutellars and supraalars always present, variously
developed; 2-4 posterior fossal and 1 parascutellar always developed. Median scutel-
lar lobe with 6 or 7 large marginal bristles and narrow, dark scales; lateral lobes with
4 or 5 large marginal bristles and with narrow scales. Paratergite bare. Pleuron usual-
ly yellowish. Bristles present on apn, ppn, ppl, stp, pra and upper mep; lower mep
bristle when present, strong, single (absent in Bisulcatus Group). Pleural scaling usu-
ally absent; a few light to dark scales on upper margin of ppn and a few light scales
along stp and between upper mep bristles. Coxae with light scales on external sur-
face; trochanters with ventral dark scales. Forefemur and midfemur dark anteriorly,
posterior side predominantly light, dark scales restricted to dorsal side; hindfemur
predominantly light on both sides, dark scaled only dorsally. Tibiae and tarsi of all
legs dark. Claws simple on all legs. Wing: Veins entirely dark scaled; plume scales re-
stricted to Rs, R,,3, R,, R;. Haltere: Stem pale, knob entirely dark scaled. Abdo-
men: Laterotergite with many moderately long bristles. Tergites II-VII with large
basolateral light patches; basal light scaling absent.
FEMALE GENITALIA (fig. 2). Only bisulcatus, the type species, studied. Seg-
ment VIII partially retracted into segment VII, apex visible, numerous bristles and
scales present; tergite VIII about 0.7 of sternite; apices truncate. Tergite IX moder-
ately broad laterally, narrow in the middle; lobe not prominent, with 4 short setae;
cowl moderately developed, finely setose, bowed laterally to join tergite IX and ar-
ticulating with sigma. Postgenital plate about 0.6 length of cercus, not joined to
cowl basally; apex rounded, with a patch of 12-14 scattered setae. Cerci short, com-
pressed, widely separated, with many short to moderately long setae; area between
cerci not lobed. Insula poorly sclerotized, with a group of 8-10 short setae, contin-
ued laterally as finely setose sigma. Spermathecae 3, one a little larger than others.
MALES (fig. 2). Coloration similar to females; sexual dimorphism of head ap-
pendages marked. Proboscis shorter to longer than forefemur. Palpus porrect; varied
in length from 0.35 to as long as the proboscis; 4 or 5 segmented, entirely dark
scaled. Whorls of flagellar segments 1-12 strongly developed, with at least 24 long
bristles; segments 12 and 13 elongate, 13 about 1.2-1.25 of 12; torus only slightly
swollen. Claws of foreleg and midleg enlarged, unequal; larger claw with a submedian
tooth; both claws with basal spicules. Hindclaws as in females. | .
MALE GENITALIA. Segment [X: Lobes of tergite [X small, approximate, usually
distinct and moundlike (indistinct in arawak) with varied number of short to moder-
ately long setae; sternite narrow, without scales or setae. Sidepiece: Roughly triangu-
lar in outline, only moderately inflated; tergal and lateral surfaces with bristles and
26 Contrib. Amer. Ent. Inst., vol. 5, no. 1, 1969
setae of varied length; sternal surface with shorter setae; scales usually absent (pres-
ent in arawak). Subapical lobe more or less distinctly differentiated into proximal
and distal divisions (poorly differentiated in Bisulcatus Group); proximal division us-
ually with 2 sclerotized rods and distal division with or without a prominent leaf.
Clasper: Relatively simple, shorter than sidepiece; either attenuate distally or distal
0.3 distinctly thickened; usually with a row of fine ridges distally on external margin
(absent in Bisulcatus Group) and 2 distinct, short subapical setae on lower surface;
spiniform subapical, short, broadened and spoon-shaped distally, apex appearing
notched in some aspects. Phallosome: Lateral plate of aedeagus with broadly sclero-
tized basal ‘‘chook’’, a posteriorly directed long apical process and a sternal spine.
Proctiger: Basolateral sclerotization strongly developed, produced into conspicuous
dorsal conical lobe below lobe of tergite IX; apex of paraproct with crown of 6-13
teeth; cercal setae small, distinct, variable in number, their alveoli usually confluent.
PUPAE. Cephalothorax: Middorsal ridge moderate. Integument uniformly lightly
to moderately pigmented. All hairs present, variously developed. Hair 5-C strongly
developed, at least 2.0 of 4-C; hair 6-C smaller than and cephalad of 7-C; hair 7-C
single; 8,9-C widely separated, 8-C single or branched (2-4) and 9-C usually single; 8-
C slightly and 9-C distinctly caudolaterad of trumpet base. Trumpet: Not placed on
tubercle; moderately long, index 8.0-12.0; moderately pigmented; tracheoid distinct,
short; apical portion not flared; pinna short. Metanotum: Hairs 10,11-C moderately
close together, removed from hair 12; hair 11-C usually single, rarely double. Abdo-
men: Integument without any distinct pattern of pigmentation. Hair 3-I single or
double, 1-II usually with at least 10 branches, faintly resembling float hair (1-1);
hair 2-III-VII not marginal but moved cephalad or cephalomesad of hair 1; hair 5-
IV-VI single or branched, at least subequal in length to its tergite (in Bisulcatus
Group, 5-IV slightly shorter than tergite length); 6-I-VI usually single; 9-II-VI cep-
halolaterad of hair 6; hair 9-VII usually triple; 9-VIII triple, at least subequal in
length to segment VIII; hair 4-VIII single or double; 1-[X apparently not developed.
Lobes on posterior margin of sternite VIII only slightly developed. Tergite VIII with
caudal lobe overlying lateral part of tergite IX. Paddle: Longer than wide; midrib
strongly differentiated; paddle margins with distinct, long hairlike spicules (absent
in Bisulcatus Group). Only 1 paddle hair (1-P) present.
LARVAE. Head: Width slightly greater than length. Labrum well differentiated
dorsally. Ocular bulge inconspicuous. Mouthbrushes numerous, filamentous. Collar
moderately well developed. Posterior tentorial pit a short distance from caudal bord-
er; maxillary suture complete, extending a short distance caudolaterad of pit. Labial
plate long. Aulaeum with distinct, filamentous spicules. Mental plate well developed,
with 7 or 8 distinct lateral teeth; central tooth not shouldered. Hair O-C distinct, re-
moved laterad of 1-C; hair 1-C strongly sclerotized, slightly curved; 2,3-C not devel-
oped; 4-C short to moderately long, either cephalomesad or cephalolaterad of 5-C;
hairs 5,6-C distinctly longer than antenna and usually 4 branched; 7-C with many
strong, barbed branches; 8-C single or double, rarely triple; 11-C strongly developed,
multiple; 13-C closer to 12 than to 11-C; hairs 14,15-C anterior; basal maxillary hair
(bmh) single or branched, when branched, spikelike or attenuate apically; 16,17-C
not developed. Antenna: Relatively short and slender, uniformly tapered; with dis-
tinct spinelike spicules in basal 0.5-0.6; pigmentation uniformly light. Hair 1-A near
middle of shaft, branched; other hairs single, short to moderately long. Thorax: Usu-
ally roughly rectangular, wider than long. Integument glabrous. Hair 1-3-P on a poor-
ly developed tubercle; 1,3-P at least moderately long, multiple; 14-P usually single
(double in bisulcatus);0-P, 1,3,14-M, 1,2-13-T stellate (strong and multiple in Schicki
Group). Abdomen: Integument of segment I glabrous; segments II-VIiI with minute
Berlin: Subgenus Micraedes of Culex a7
spinelike spicules laterally. Hairs 1,2,4,11,13-I, 1,2,5,9,13-II, 1,2,5,7,9,13-IlI-V1, 1,
2,5,8,10,13-VII usually stellate (strong and multiple in Schicki Group); hair 2 always
in the anterior part of tergite; 6-I long, at least double; 6-II single; 7-I long, double;
7-II at least double; 6-III-VI shorter, double, branches subequal. Segment VIII:
Comb in a patch of 3 or 4 irregular rows; comb scales slender, simple, not spatulate.
Hairs 1,5-VIII multiple, sometimes stellate; 1,2-VIII not on sclerotized tubercles or
plates. Siphon: Long, index ranging from 9.0 to 20.0. Acus distinct, attached, with
ventral projection. Pecten terminating in basal 0.4. Siphonal hairs strongly devel-
oped, 6 pairs; 4 pairs subventral, progressively shorter caudad, proximal distinctly
caudad of last pecten tooth; 2 pairs subdorsal, usually branched. Subapical hair (2-S)
slightly recurved and with small subbasal branch. Anal Segment: Saddle complete;
‘integument imbricate or spiculate; caudal margin with long spines, some branched
apically; acus not developed. Hair 1-X short to moderately long, single, double or
forked; 2-X long, at least double, without subbasal branches; 3-X long, single. Ven-
tral brush (4-X) with 5 pairs of 2-4 branched hairs, all on grid with distinct lateral
bar not attached to saddle. Gills varied in length, ventral subequal to dorsal.
KEYS TO GROUPS AND SPECIES
ADULTS
Bisulcatus Group
i Lower mep area without bristles; palpus of both sexes slender, about 0.35 of
proboscis length . . . . 1. bisulcatus; 2. antillummagnorum; 3. arawak
Lower mep area with | strong bristle; palpus of both sexes moderately thick,
female less than 0.28 and male about 0.9 of proboscislength. . . . .2
Schicki and Erethyzonfer Groups
2(1). Scales on upper margin of ppn white; lower mep area dark brown (Schicki
CN ge os . . 4. schicki; 5. sandrae
Scales on upper margin of ppn brown to blackish: lower mep area yellowish
like rest of pleuron (Erethyzonfer Group) . . .. . . 6.erethyzonfer
MALE GENITALIA
(5. sandrae unknown)
i, Subapical lobe not clearly differentiated into divisions, distal division appear-
ing as distal sternal appendage of proximal; subapical lobe without prom-
inent leaf (Bisulcatus Group) . . a
Subapical lobe differentiated into proximal and distal divisions: “subapical
Te EE OE RN ee ny tate cit cma eae
Bisulcatus Group
2(1). Paraproct with only 6 or 7 apical teeth; clasper with a moderately long sub-
DASH SOLA ON CRLOENAL WATER NN ae i Bo eee
28
3(2).
4(1).
2(1).
3(1).
4(3).
Contrib. Amer. Ent. Inst., vol. 5, no. 1, 1969
Paraproct with at least 10(10- 13) ae teeth; i without any subbasal
setae on external margin... . aw He ee
Ninth tergite lobe usually with 3(2-5) short weak setae; 1 bristle on base of
tergal surface of proximal division of subapical lobe distinctly sclerotized
and saberlike . . . . . . IL. bisulcatus
Ninth tergite lobe usually with 6(6-9) ioderately long setae; none of the
bristles on base of tergal surface of proximal division sclerotized and saber-
ik ee be oe ee
Schicki and Erethyzonfer Groups
Proximal division of subapical lobe with 2 subequal rodlike appendages;
ninth tergite lobe with at least 10 moderately long setae . :
6. erethyzonfer
Proximal division of subapical lobe with only 1 ‘distinct rodlike appendage;
ninth tergite lobe with 3-6 short, weak setae . . . .. . . .4.schicki
PUPAE
(3. arawak unknown)
Paddle margins smooth; hair 9-VIII very long, about 2.0 length of tergite
VIII (Bisulcatus Group) . . 2
Paddle margins with long hairlike spicules: hair 9-VIII moderately Jone, sub-
equal in length to tergite VIII . Pe es 3
Bisulcatus Group
Hair 6-I,I] moderately long, usually subequal to or i i longer than 7-I,II
1. bisulcatus
Hair é I Il long, usually about 2. 0 length of a4 I } : os antillummagnorum
Schicki and Erethyzonfer Groups
Hair 6-IV,V long, exceeding apex of succeeding segment. . . . .4. schicki
Hair 6-IV,V shorter, barely extending to middle of succeeding segment . .4
Hair 10-C triple: 1-II usually 4 branched . . . . 5. sandrae
- Hair 10-C ss 7 branched a ay hair 1-Il at least 10 branched .
6. erethyzonfer
LARVAE
Hair 8-C single; 4-C usually single and cephalomesad of 5-C; basal maxillary
hair (bmh) strong, spikelike and at least double (Bisulcatus Group). . .2
Hair 8-C double or triple; 4-C at least double and cephalolaterad of 5-C; bas-
sal maxillary hair (bmh) weak and single; if branched, then with attenu-
Bted Bianices ee er ee ee ee ee
Berlin: Subgenus Micraedes of Culex 29
Bisulcatus Group
2(1). Metathoracic hair 1-T with only 3 or 4 branches; 2-II single or double
ge G2 Te a A es ey . . .d. arawak
- Metathoracic hair 1-T with at least 15 branches; 24 stellate, with at least 15
Branches io, UA Rae aes a a ea 2 eee Foe ee
3(2). Hair 6-I with at least 5(5 or 6) branches . . . .. . . .. . I. bisulcatus
Hair 6-I usually with 3 branches . . . . .. . . . 2. antillummagnorum
Erethyzonfer Group
4(1). Saddle integument distinctly spiculate at 100X; abdominal hairs 1 ,2,5,13-II-
VI stellate, with at least 15 strong spikelike branches; caudal margin of
saddle with spines extending ventrad of hair 1-X. . . . 6.erethyzonfer
Saddle integument imbricate, without spicules visible at 100X; abdominal
hairs 1,2,5,13-IJ-VI multiple, usually with less than 12 attenuated branch-
es; caudal margin of saddle without spines ventrad of hair 1-X ... .5
Schicki Group
5(4). Hair 2-X usually double, rarely triple; proximal subdorsal siphonal hair short,
weakly developed and about 0.25 of hair 1-S . . . . «4. schicki
Hair 2-X at least 4 branched (4-6); proximal subdorsal siphonal hair long,
strongly developed and subequal in length to hair 1-S. . . . 5.sandrae
BISULCATUS GROUP
FEMALES. Small to moderate in size. Head: Erect scales usually yellowish to
pale brown. Palpus moderately long, slender, about 0.35 to 0.38 of proboscis length;
segment 4 about 2.0 of 3. Thorax: Mesonotal scales auburn. Pleural integument yel-
lowish. Upper marginal scales on ppn auburn. Lower mep area without any bristles.
Abdomen: Sternites II-VII predominantly with creamy scales, occasionally a few
apical dark ones on distal segments.
MALES. Similar to female in coloration. Palpus narrower than in female, about
0.35-0.38 of proboscis length and 5 segmented; segment 4 about 1.5-1.6 of 3; seg-
ment 5 minute.
MALE GENITALIA. Ninth tergite lobe small, usually moundlike (indistinct in
arawak), with 2-9 short to moderately long setae. Sidepiece roughly conical, length
2.0-2.5 of greatest width; tergal surface with bristles and setae and usually without
scales (present in arawak); sternal surface with short to moderately long setae. Sub-
apical lobe situated at about middle, directed mesad and not clearly differentiated
into divisions, the distal division appearing as distal sternal appendage of proximal;
proximal division prominent, split apically, each part bearing a sclerotized rod; distal
division with relatively simple setae; leaf absent. Clasper about 0.6-0.8 of sidepiece
length, relatively simple, slender; distal 0.4 with 2 subapical setae on lower surface.
Apical process of lateral plate long, truncate from dorsal aspect; sternal spine direct-
ed laterad. Paraproct with at least 10(10-13) apical teeth (only 6 or 7 in arawak);
cercal setae distinct, 2-5 in number.
30 Contrib. Amer. Ent. Inst., vol. 5, no. 1, 1969
PUPAE. Cephalothorax: Hair 4-C short, usually double; 5-C distinctly longer than
distance between its base and that of trumpet; 10-C at least 4 branched. Abdomen:
Hair 1-II with many branches (25 or more), faintly resembling float hair; 1-III-VII
usually single or double; 2-II-VI about 2.0 of 9-II-VI; hair 2-VII always within basal
0.66 of its tergite; 5-V,VI at least 1.5 of succeeding segment; 9-III-VII distinctly cep-
halad of hair 6; hair 9-VII distinctly longer than 6-VII, extending beyond apex of
segment; 9-VIII at least 2.0 length of segment VIII. Paddle: Elongate, apex slightly
produced; margin smooth, without any indication of spicules.
LARVAE. Head: Hair 4-C short, usually single and slightly cephalomesad of 5-C;
hairs 8,10-C always single; basal maxillary hair (bmh) with 2-4 strong, spikelike
branches. Antenna: Hair 1-A at about 0.5 from base. Thorax: Roughly rectangular
in outline, wider than long. Hair 4-P at least triple; 9-M double or triple. Abdomen:
Hairs 1,2,4,11,13-1, 1,2,5,9,13-I1, 1,2,5,7,9,13-101-VI, 1,2,5,8,10,13-VII distinctly
stellate, with many spikelike branches; 2-I-VI distinctly cephalad of 1 (slightly cep-
halolaterad in arawak). Segment VIII: Individual scale usually terminating in a med-
ian spine (absent in arawak). Hairs 1,5-VIII usually stellate. Siphon: Long, index
about 10.0-20.0. Pecten terminating before middle. Subventral hairs moderately
long, progressively shorter distally; proximal subdorsal hair basad of hair 1-S and us-
ually within last pecten tooth. Anal Segment: Caudolateral border of saddle with
very long marginal spines, some with lateral fringes; no distinct spines ventrad of hair
1-X. Hair 1-X moderately long, single. Saddle body with rows of minute hairlike
spicules. Hair 2-X usually triple. Gills subequal, less than 1.5 of dorsal saddle length.
DISCUSSION. The Bisulcatus Group is strongly differentiated from the other
groups of the subgenus. Both sexes of the adults have a narrow palpus which is only
0.35-0.38 of the proboscis length. The male genitalia are readily recognized by the
position of the subapical lobe of sidepiece and the absence of a row of fine ridges on
distal 0.4 of clasper. The larvae are immediately separated by the position of hair 4-
C and the spikelike basal maxillary hair. The pupae are easily differentiated by the
absence of marginal spicules on the paddle.
The distribution of the Bisulcatus Group is in general similar to but more restrict-
ed than that of the Walkeri Section of the subgenus Howardina of Aedes (Berlin,
1969). The group is known at present only from islands in the Greater and Lesser
Antilles. In the present study I am recognizing 3 species in the group on the basis of
correlated features of the adults and the immature stages. All 3 species have been
mistaken for americanus in the past. C. bisulcatus, the type species of the subgenus,
is definitely known only from the Leeward and Windward groups of the Lesser An-
tilles, having been reported from St. Croix southward to St. Lucia; antillummagnor-
um has been collected in Cuba, Hispaniola, Puerto Rico and St. Thomas; and arawak
is endemic to the island of Jamaica (fig. 1).
The immature stages of the Bisulcatus Group have been collected primarily in epi-
phytic and terrestrial bromeliads.
1. Culex (Micraedes) bisulcatus (Coquillett)
Figs. 1-4
1906. Micraedes bisulcatus Coquillett, 1906a:185-186. TYPE: Lectotype 6 (82.3) with associ-
ated larval and pupal skins and genitalia slide (1648), La Soufriere, Guadeloupe, elev.
3,000 ft, 30 July 1905, A. Busck [USNM, 8291; designation of Stone and Knight 1957:
44].
Berlin: Subgenus Micraedes of Culex 31
Culex (Micraedes) bisulcatus of Dyar (1918:90,102; 1922:96; 1923:188); Bonne and Bonne-Wep-
ster (1925:275); Floch and Abonnenc (1945 :36-38); Belkin (1968:11).
Culex (Aedinus) bisulcatus in part of Stone (1967:218; 1969:7).
Culex bisulcatus of Dyar and Knab (1906:205,208); Howard, Dyar and Knab (1915:306-308, in
part). ,
Micraedes bisulcatus of Coquillett (1906b:24; 1910:569); Theobald (1910:486).
Culex (Micraedes) americanus in part of Edwards (1932:219); Lane (1939:76).
Culex (Aedinus) americanus in part of Stone, Knight and Starcke (1959:281); Belkin, Schick and
Heinemann (1965 :27); Forattini (1965 :34); Bram (1967:17).
Culex (Tinolestes) americanus in part of Lane (1953:389-390); Duret and Damasceno (1955:395).
Culex (Melanoconion) americanus in part of Dyar (1928:343-344).
Culex americanus of van der Kuyp (1948:748,749; 1953:146; 1954:58).
FEMALE (fig. 2). Wing: 2.8 mm. Proboscis: 1.7 mm. Forefemur: 1.6-1.7 mm.
Abdomen: 2.1 mm. As described for the subgenus and the group, with the following
additional features. Decumbent scales on dorsum of head narrow, creamy. Proboscis
length variable; subequal to forefemur in Nevis, Montserrat, Guadeloupe, Dominica
and Martinique populations; slightly longer in St. Kitts, Antigua and St. Lucia popu-
lations. Mesonotal scales auburn, linear. Antealer area above paratergite with a few
scattered auburn scales. Basolateral light patches on abdominal tergites creamy.
MALE (fig. 2). Wing: 2.5 mm. Proboscis: 1.6 mm. Forefemur: 1.5-1.6mm. Slight-
ly smaller in size than female and similar in coloration. Palpus length as in female.
Proboscis subequal to forefemur in Nevis, Montserrat, Guadeloupe and Martinque
and longer than forefemur in St. Kitts, Antigua, Dominica and St. Lucia populations.
MALE GENITALIA (fig. 3). Diagnostic characters as in the key. Readily separ-
ated from antillummagnorum by the presence of usually 3(2-5) short, weak setae on
ninth tergite lobe; from arawak by the presence of at least 10 apical teeth on para-
proct. Ninth tergite lobe prominent, usually with 3(2-5) short setae. Sidepiece length
about 2.0 of greatest width; tergal and lateral surfaces with many long bristles, stern-
al surface with short setae; no scales evident. Proximal division of subapical lobe
prominent, split apically into 2 unequal parts, tergal part bearing a smaller and stern-
al a larger saber; a smaller narrow saber arising from base of tergal surface; distal di-
vision with 8-10 relatively simple setae, of which 4 or 5 lanceolate, 1 longer seta
with attenuated apex and 1 with a recurved apical barb. Clasper about 0.8 of side-
piece length, relatively simple, slender. Paraproct with 10-13 apical teeth; cercal set-
ae 2-5.
PUPA (fig. 3). Abdomen: 2.6 mm. Trumpet: 0.4 mm; index about 8.0. Paddle:
0.75 mm. As figured; diagnostic characters as in the key. Readily separated from
antillummagnorum by hair 6-I,II either subequal to or slightly longer than 7-I,II.
Cephalothorax: Integument lightly pigmented. Hair 5-C double or triple; 6-9-C usu-
ally single; 11-C single and 12-C triple (2 or 3). Abdomen: Integument lightly pig-
mented, progressively lighter caudad. Hair 1-II with at least 30 branches; 6-I,II single,
moderately long, either subequal to or slightly longer than 7-I,II; hair 5-V single; 5-
VI double; 6-III-VI usually single; 6-VII usually triple (2 or 3), distinctly shorter
than 9-VII; hair 9-VIII triple. Paddle: Elongate, longer than wide, at least 2.0 of seg-
ment VIII. Male genital lobe extending to 0.43 and female barely to 0.2 of paddle.
LARVA (fig. 4). Head: 0.95 mm. Siphon: 1.4-1.5 mm. Anal Saddle: 0.25 mm.
As figured; diagnostic characters as in the key. Readily separated from antillummag-
norum by the 5 or 6 branched hair 6 on segment I; from arawak by hair 14-P usually
double and hair 2-II with at least 15 stellate branches. Head: Width about 1.05 of
length. Hair 4-C usually single; 7-C usually with 10(9-11) and 9-C usually with 4(4-7)
32 Contrib. Amer. Ent. Inst., vol. 5, no. 1, 1969
branches; 15-C short, double, not extending to base of mental plate. Mental plate
with a strong median tooth and 7 or 8 distinct teeth on each side. Antenna: Length
about 0.45 of head, with distinct spicules in basal 0.5; all hairs single except 1(9,7-
11). Thorax: Hair O-P usually with 23(22-24) stellate branches; 7-P single; 14-P
spikelike, double or triple, rarely single; 1-M usually with 25(23-26) branches. Abdo-
men: Stellate hairs usually with at least 15 branches; 6-I with 5 or 6 branches; 2-II
with at least 15 stellate branches; 2-I-VI distinctly cephalad of hair 1. Segment VIIT:
Comb scales in a patch of 3 or 4 irregular rows, about 55-67 in number; scales in
proximal row fringed to apex; distal scales with a recurved apical spine. Siphon: In-
dex 14.0-16.0. Integument lightly pigmented, faintly imbricate in basal 0.1-0.2; sub-
ventral hairs moderately long, usually with less than 5 branches, progressively small-
er caudad; subdorsal hairs moderately long, double. Pecten extending to 0.35 of
siphon; individual tooth long, with minute denticles on ventral border but not ex-
tending to apex. Anal Segment: Integument lightly pigmented, weakly spiculate;
caudolateral border with long spines, some branched apically. Hair 1-X single; 2-X
triple. Gills narrow, moderately long, about 1.4-1.5 of dorsal saddle length.
SYSTEMATICS. The incorrect synonymy of bisulcatus with americanus (Neveu-
Lemaire, 1902) by Dyar (1928:344) was questioned by Floch and Abonnenc (1945:
37) and Fauran (1961:45) but persisted until Stone (1967:218) treated americanus
as a nomen dubium and restored bisulcatus to specific status, as was also done inde-
pendently by Belkin (1968:11). enue
The present diagnosis of bisulcatus in the restricted sense is based primarily on
topotypic material from Guadeloupe. Populations from other islands in the Leeward
and Windward groups of the Lesser Antilles agree in general with the Guadeloupe
population in all stages except for the number of bristles on the ninth tergite lobe of
the male genitalia which tend to be more numerous in the northernmost popula-
tions, approaching the condition in antillummagnorum. The only significant varia-
tion found in the different insular populations is a non-clinal one in the length of the
proboscis. The proboscis is subequal to the forefemur in both sexes from Nevis,
Montserrat, Guadeloupe and Martinique; longer in both sexes from St. Kitts, Antig-
ua and St. Lucia; longer in the males and subequal in the females from Dominica.
The record of bisulcatus from the southern Virgin Islands (St. Croix) is based on
the genitalia of a single male [USNM] ; these agree in general with the other popula-
tions of bisulcatus and not with antillummagnorum.
BIONOMICS. The immature stages of bisulcatus have been collected in the leaf
axials of arboreal and terrestrial bromeliads at low (50 ft) to moderate (3200 ft) ele-
vations. Several adults, including 2 blooded females resting on tree trunks, were tak-
en in a sweeping collection at 1500 hours on Guadeloupe (FWI 195).
DISTRIBUTION (fig. 1). Islands of the Leeward and northern Windward groups
of the Lesser Antilles from Sint Maarten to St. Lucia; St. Croix in the southern Vir-
gin Islands. Not known from the southern Windward islands (St. Vincent, Grena-
dines, Grenada and Barbados). Material examined: 1214 specimens; 113 6, 88 2, 237
pupae, 776 larvae; 191 individual rearings (35 pupal, 93 larval, 63 incomplete).
ANTIGUA. St. John: Table Hill, elev. 175 ft, 25 Sept 1965, R. Martinez and A. Guerra (ANT
103), 7 L [UCLA]. St. Mark: Boggy Peak, elev. 1300 ft, 22 Sept 1965, R. Martinez and A. Guerra
(ANT 82), 2 L [UCLA]. St. Mary: Walling’s Dam, elev. 300-500 ft, 15 Sept 1965, T.H.G. Aitken,
R. Martinez and A. Guerra (ANT 45,48,50), 4 lpt (45-12,14-16), 2 IP (45:10,13), 1 6, 5:9, 6 P,
21 L (45), 2 IP (48-10,11), 4 6, 3 9, 7 p, 23 L (48), 1 IP (50-10), 19 L (50); 27 Sept 1965, R. Mar-
tinez and A. Guerra (ANT 112,113), 2 Ipd (112-11,12), 1 Ip? (112-17), 1 pd (112-100), 2 Ip (112-
10,15), 2 IP (112-13,16), 3 L (112), 1 Ip? (113-13), 1 pd (113-100), 1 1d (113-108), 1 lp (113-12),
1 IP (113-11), 1 P, 9 L [UCLA]. St. Paul: Liberta, 23 Sept 1965, R. Martinez and A. Guerra
Berlin: Subgenus Micraedes of Culex 33
(ANT 94), 2 P, 5 L, 3 1 [UCLA]. Sugar Loaf Mountain, elev. 750-800 ft, 26 Sept 1965, R. Mar-
tinez and A. Guerra (ANT 106-110), 1 pd (106-100), 10 L (106), 2 IP (107-10,11), 8 L (107),
4 IP (108-10-13), 2 9, 2 p, 5 L (108), 1 IP (109-12), 2 L (109), 1 IP (110-10), 8 L(110) [UCLA].
DOMINICA. St. David: Crete Palmiste, elev. 1400 ft, 18 June 1965, R. Martinez and A. Guerra
(DOM 120), 1 Ipd (120-11), 6 L [UCLA]. Fond Figues, elev. 400-500 ft, 24 June 1965,
R. Martinez and A. Guerra (DOM 146), 2 pd (146-100,101), 9 L [UCLA]; 30 Jan 1965, W.W.
Wirth, 1 2 (66-29); 9 Feb 1965, W.W. Wirth, 1 ¢ (66-16), 1d, 1 9; 12 Apr 1966, R.J. Gagne, 1 9;
29 Apr 1966, R.J. Gagne, 1 ¢ (67-41) [USNM]. Grand Fond, elev. 300 ft, 18 June 1965, R. Mar-
tinez and A. Guerra (DOM 113,114), 5 L (113), 1 lpd (114-24), 1 IP (114-20), 8 L [UCLA]. Rosa-
lie, Bois Diable Ridge, elev. 1400 ft, 11 June 1965, R. Martinez and A. Guerra (DOM 72), 2 IP
(72-10,11), 1 L [UCLA]. Junction of roads to Rosalie and Castle Bruce, 29 Mar 1966, R.J. Gagne,
1 6 (67-39), 1 9; 23 Apr 1966, R.J. Gagne, 1 3 (67-36), 2 9? [USNM]. St. George: Morne Anglais,
elev. 3000 ft, 25 June 1965, R. Martinez (DOM 147-149,152), 1 Ipd (147-20), 2 L (147), 1 L
(148), 1 pd (149-100), 2 p, 1 L (149), 1 Ipd (152-10) [UCLA]. Roseau, elev. 50 ft, 5 June 1965,
T.H.G. Aitken, R. Martinez and A. Guerra (DOM 12), 1 lpd (12-21), 1 IP (12-20) [UCLA]. Roseau,
L’Etang, elev. 2600 ft, 8 June 1965, T.H.G. Aitken, R. Martinez and A. Guerra (DOM 44,46,51,
53), 1 IP (44-30), 1 lpd (46-10), 1 IP (46-11), 3 L (46), 1 lp? (51-10), 1 p? (51-100), 4 L (51), 1 pd
(53-100), 2 L (53) [UCLA]. Tan Cred, elev. 2000 ft, 9 June 1965, R. Martinez and A. Guerra
(DOM 56), 2 IP (56-10,11), 10 L [UCLA]. St. John: Morne Diablotin, elev. 3000 ft, 24 June
1965, A. Guerra (DOM 161,162), 2 IP (161-10,13), 7 L (161), 1 Ip? (162-11), 1 p? (162-100),
2 L (162) [UCLA]. Syndicate Estate, elev. 2000 ft, 24 June 1965, A. Guerra (DOM 160),
1 Ipd (160-10) [UCLA]. St. Joseph: Clarke Hall, 20 Feb 1964, D.F. Bray, 1 6 (67-44) [US-
NM]. Layon Park Estate, elev. 600 ft, 24 June 1965, R. Martinez (DOM 142), 1 IP (142-10),
4 L [UCLA]. Layou Valley, 18 Apr 1959, R.H. Darsie, 1 Ip? (72-2), 2 pd (72-3,5A) [USNM].
Mero, elev. 200 ft, 14 June 1965, R. Martinez and A. Guerra (DOM 86), 1 pd (86-100), 1 IP (86-
10), 2 L [UCLA]. St. Luke: Champigny Estate, elev. 600-1000 ft, 6 June 1965, T.H.G. Aitken,
R. Martinez and A. Guerra (DOM 14,24), 2 lpd (14-10,11), 4 3, 2 2, 6 p, 3 L(14), 2 IP (24-10,11),
2 P,3 L (24) [UCLA]. South Chiltern, elev. 600-800 ft, 6 June 1965, T.H.G. Aitken, R. Martinez
and A. Guerra (DOM 25-27), 1 Ipd (25-10), 1 IP (25-11), 3 L (25), 1 Ip? (26-11), 1 p? (26-100),
1 IP (26-10), 8 L (26), 1 p? (27-100), 2 L (27) [UCLA]. St. Patrick: Geneva Estate, elev. 300 ft,
10 June 1965, R. Martinez and A. Guerra (DOM 64), 1 Ipd (64-10), 1 pd (64-100), 1 IP (64-11),
6 L [UCLA]. St. Paul: Brigandin, elev. 2000 ft, 22 June 1965, R. Martinez and A. Guerra (DOM
125), 1 p? (125-100), 5 L [UCLA]. Pont Casse, elev. 2200-3000 ft, 22 June 1965, R. Martinez
and A.Guerra (DOM 126,128,129), 1 p? (126-100), 8 L (126), 5 L (128), 2 P, 5 L (129) [UCLA];
12 Feb 1965, W.W. Wirth, 1 6 (66-34); 25 Feb 1965, W.W. Wirth, 1 6 (66-33); 10 Apr 1966, R.J.
Gagne, 1 2; 11 Apr 1966, R.J. Gagne, 1 6 (67-40); 5 May 1966, R.J. Gagne, 1 9; 19 Apr 1959,
R.H. Darsie, 1 IP (73-6), 1 P (73-3), 2 L (73), 3 1(73-1,2,8), 1 Ipé (74-1), 1 IP (75-3), 1 lp (76-7),
1 3 (78-3A), 1 lp (78-3), 1 p (78-1), 1 L. (78-4) [USNM] . Parish not specified: Manets Gutter, 10
Mar 1965, W.W. Wirth, 1 9 [USNM].
GUADELOUPE. Basse-Terre: Bains Jaunes, elev. 3000 ft, 21 Sept 1965, P. Fauran (FWI 248),
1 Ip? (248-10) [UCLA]. La Soufriere, elev. 3000 ft, July 1905, A. Busck, 1 1d (82.1) [USNM].
Lamentin, Duclos, elev. 500 ft, 19 Jan 1965, P. Fauran (FWI 186-191), 1 lpd (186-100), 3 Ipd
(187-101 103,104), 2 Ip? (187-100,102), 4 L (187), 3 Ipd (188-100,101,107), 3 lp? (188-104,106,
108), 1 1d (188-105), 2 Ip (188-102,103), 6 L (188), 6 Ipd (189-100,103,105-107,110), 1 Ip? (189-
104), 1 Ip (189-102), 2 IP (189-111,112), 2 Ipd (190-101 ,103), 1 Ip (190-102), 1 IP (190-100), 1
Ip? (191-100) [UCLA] . Route de Traversee, elev. 680-1200 ft, 18 Sept 1965, P. Fauran (FWI 241-
244), 1 L (241), 1 lpd (242-13), 6 Ip? (242-10-12,14-16), 2 pd (242-100,101), 3 p, 18 L, 31
(242), 1 Ipd (243-10), 1 Ip? (243-11), 2 pd (243-102,103), 2 p? (243-100,101), 1 6, 1 p, 31(243),
2 Ipd (244-11,12), 2 Ip? (244-10,13), 3 L (244); 24 Sept 1965, P. Fauran (FWI 250,251), 1 Ip?
(250-10), 5 L (250), 1 Ips (251-11), 2 Ip? (251-10,12), 1 pd (251-100), 1 19 (251-102), 1 P, 20 L,
41(251) [UCLA] . Grande-Terre: Pointe-a-Pitre, Joliviere, adult collection, 26 Jan 1965, P. Fauran
(FWI 195), 13 6,79 [UCLA].
MARTINIQUE. Fort-de-France, Route de la Trace, elev. 1860 ft, 16 J uly 1965, P. Fauran (FWI
ce Ipd (M 3-10), 2 Ip? (M 3-11,12),1 3, 1 L(M 3), 1 Ipod (M 4-10), 1 Ip? (M 4-11), 1 p(M
34 Contrib. Amer. Ent. Inst., vol. 5, no. 1, 1969
MONTSERRAT. St. Anthony: Chance Mountain, elev. 2800-2850 ft, 5 Oct 1966, T.H.G. Ait-
ken, R. Martinez and A. Guerra (MNT 30-33), 1 Ip (30-22), 2 IP (30-21,23), 12 L (30), 2 Ipd
(31-11,12), 1 lp? (31-10), 1 IP (31-13), 9 L (31), 15 L (32), 1 Ipd (33-10), 2 Ip? (33-11,12), 2 IP
(33-13,14), 11 L (33) [UCLA]. 0.5 mi SE of Galways Soufriere, elev. 1750 ft, 12 Oct 1966, R.
Martinez and A. Guerra (MNT 59), 17 L [UCLA]. Lower Gages Soufriere, elev. 1250 ft, 6 Oct
1966, T.H.G. Aitken, R. Martinez and A. Guerra (MNT 37), 1 lp? (37-10), 9 L [UCLA] . O’Gara’s
Estate, elev. 100 ft, 11 Oct 1966, R. Martinez and A. Guerra (MNT 56), 6 L [UCLA] . Richmond
Estate, elev. 50 ft, 10 Oct 1966, R. Martinez and A. Guerra (MNT 47), 24 L [UCLA] . Waterworks
Estate, elev. 300 ft, 18 Oct 1966, R. Martinez and A. Guerra (MNT 123), 1 pd (123-100), 1 d,
1 p, 20 L, 11 [UCLA]. St. Peter: Fogarty Village, elev. 500 ft, 17 Oct 1966, R. Martinez and A.
Guerra (MNT 114), 12 L [UCLA]. Frith Village, 4 Oct 1966, T.H.G. Aitken, R. Martinez and
A. Guerra (MNT 23), 2 lp? (23-10,11), 1 pd (23-100), 1 p? (23-101), 26 L [UCLA] . Plymouth,
0.5 mi NE of Belham Bridge, elev. 250 ft, 4 Oct 1965, T.H.G. Aitken, R. Martinez and A. Guerra
(MNT 21), 1 IP (21-10), 2 L [UCLA]. Woodlands Estate, elev. 300 ft, 17 Oct 1966, R. Martinez
and A. Guerra (MNT 115,116), 1 lp? (115-10), 2 6,2 p, 14 L(115), 6 L (116) [UCLA].
NEVIS. St. George: Nevis Peak, elev. 2600-3200 ft, 24 Aug 1966, R. Martinez (NEV 19-22),
1 IP (19-10), 1 P, 5 L(19), 1 lpé (20-10), 1 IP (20-11), 2 L (20), 3 L (21), 1 pd (22-100), 3 L (22);
8 Sept 1966, R. Martinez and A. Guerra (NEV 39-42), 1 Ipd (39-12), 1 Ip? (39-11), 1 pd (39-
100), 4 L (39), 1 Ip? (40-11), 8 L (40), 1 Ipd (41-11), 7 L (41), 1 Ipd (42-11), 1 pd (42-100),
5 L (42) [UCLA].
SABA. Peak Hill, Rendez Vous, and Upper Hellsgate, Apr 1947 (van der Kuyp 1954:91).
SINT EUSTATIUS. Locality and date not specified, E. van der Kuyp, 1 L [USNM]. De Kant
of the Quill, 12 July 1949, 23 L; Big Gut near base of White Hall, 6 July 1949, 4 L; near top of
White Hall, 6 July 1949, 1 L (van der Kuyp 1953:146).
SINT MAARTEN. Marigot Hill, 7 Oct 1947, E. van der Kuyp, 2 6 [USNM]. Locality and date
not specified, E. van der Kuyp, 3 L [UCLA, USNM]. Base of Meschrine Hill, S of Simon Bay
bridge, 27 May 1949, 2 P, 17 L(van der Kuyp 1953:146).
ST. CROIX. Crique, Apr 1938, H.A. Beatty, 1 d genitalia [USNM].
ST. KITTS. St. John: Slopes of Mt. Misery, elev. 2900 ft, 17 Aug 1966, T.H.G. Aitken, R.
Martinez and A. Guerra (KIT 28), 1 Ipod (28-21), 1 lp? (28-11), 1 IP (28-12), 12 L [UCLA]. Lower
Mt. Misery, elev. 2900 ft, 17 Aug 1966, T.H.G. Aitken, R. Martinez and A. Guerra (KIT 29,30),
1 Ip? (29-11), 1 P, 3 L (29), 1 lpd (30-10), 1 p? (30-100), 1 IP (30-11), 6 L (30) [UCLA]. Sz.
Thomas: Wingfield Manor Estate, elev. 500 ft, 28 Aug 1966, A. Guerra (KIT 68), 1 L [UCLA].
ST. LUCIA. Barre de L’Isle, elev. 910 ft, 29 July 1964, R. Martinez and A. Guerra (LU 110),
3 L [UCLA]. Black Bay, elev. 300 ft, 29 July 1964, R. Martinez and A. Guerra (LU 120), 3 IP
(120-10-12), 6 L [UCLA]. Canaries, elev. 200 ft, 22 July 1964, R. Martinez and A. Guerra (LU
77), 1 p? (77-100), 3 L [UCLA]. Chantin, elev. 600 ft, 31 July 1964, R. Martinez and A. Guerra
(LU 136-139), 1 lpé (136-10), 4 p, 6 L (136), 3 6, 1 9,4 L (137), 2 L (138), 1 Ip? (139-10), 5 L
(139) [UCLA]. Micoud, Mahaut, elev. 650 ft, 27 July 1964, A. Guerra (LU 100), 1 IP (100-20),
7 L [UCLA]. Reunion, elev. 275 ft, 24 July 1964, R. Martinez and A. Guerra (LU 87,88), 1 pd
(87-101), 5 L (87), 1 IP (88-20), 1 6, 6 L (88) [UCLA] . Soufriere, elev. 450 ft, 22 July 1964, R.
Martinez and A. Guerra (LU 79), 1 lp? (79-20), 1 IP (79-21), 1 L [UCLA].
2. Culex (Micraedes) antillummagnorum Dyar
Figs. 1,5,6
1928. Culex (Melanoconion) antillummagnorum Dyar, 1928:344. TYPE: Lectotype ¢ (416)
with genitalia slide (785), San Antonio de los Banos (La Habana), Cuba, date not speci-
fied, J.H. Pazos [USNM, 40778; designation of Stone and Knight 1957:43].
Culex (Micraedes) antillummagnorum of Edwards (1932:219); Lane (1939:76); Belkin (1968:11).
Culex (Melanoconion) antillummagnorum of Gerry (1932:44-45 49); Komp (1935:9).
Berlin: Subgenus Micraedes of Culex 35
Culex (Aedinus) bisulcatus in part of Porter (1967:40); Stone (1967:218; 1969:7).
Culex bisulcatus of Howard, Dyar and Knab (1915:306-308, in part); Root (1922:395; 1927:
464).
Culex (Aedinus) americanus in part of Stone, Knight and Starcke (1959:281)- Belkin, Schick and
Heinemann (1965:16).
Culex (Melanoconion) americanus of Tulloch (1937:148,157); Montchadsky and Garcia (1966:
44-45 84).
Culex (Melanoconion) americanus in part of Rozeboom and Komp (1955:77).
Culex (Tinolestes) americanus in part of Lane (1953:389-390).
Culex americanus of Fox (1953:179); Maldonado-Capriles, Pippin and Kuns (1958:68).
FEMALE. Wing: 2.9 mm. Proboscis: 1.7 mm. Forefemur: 1.7 mm. Abdomen:
1.6-1.7 mm. Similar to bisulcatus in general characters. Proboscis subequal in length
to forefemur.
MALE. Wing: 2.5 mm. Proboscis: 1.6 mm. Forefemur: 1.45 mm. Similar to fe-
male, differing only in the length of proboscis which is longer than forefemur.
MALE GENITALIA (fig. 5). Diagnostic characters as in the key. Readily separated
from bisulcatus by the presence of at least 6 moderately long setae on ninth tergite
lobe; from arawak by the presence of at least 11 apical teeth on proctiger. Generally
similar to bisulcatus, differing in the following. Ninth tergite lobe prominent, with 6-
9 moderately long setae. Sidepiece length about 2.5 of greatest width, with fewer
long bristles on tergal and lateral surfaces; sternal surface with short setae; no scales
evident. Proximal division of subapical lobe prominent, split apically into 2 unequal
parts, tergal bearing a narrower and sternal a broader rod with recurved apex; a
small narrow seta arising from base of tergal surface; distal division with 5 or 6 rela-
tively simple setae, 1 longer and with recurved apex and 1 with a recurved apical
barb. Clasper about 0.75 of sidepiece length, relatively simple. Paraproct with 11 or
12 apical teeth; cercal setae 2 or 3.
PUPA (fig. 5). Abdomen: 2.8 mm. Trumpet: 0.45 mm; index about 9.0. Paddle:
0.7 mm. As figured, diagnostic characters as in the key. Description based on a single
skin from Puerto Rico. Readily separated from bisulcatus by very long hair 6-I,II, at
least 2.0 of 7-I,II. Cephalothorax: Integument lightly pigmented. Hair 5-C triple; 6,
9-C single; 7,8-C double; 11-C single, 12-C double. Abdomen: Integument lightly pig-
mented, progressively lighter caudad. Hair 1-II with at least 15 branches (15-25);
hair 6-I,II single, long, at least 2.0 of hair 7-I,I]; hair 5-V single, 5-VI double; 6-[HI-VI
double; 6-VII with 2-4 branches, about 0.5 of 9-VII; hair 9-VIII triple. Paddle:
Elongate, longer than wide, about 2.0 of segment VIII. Female genital lobe barely
extending to 0.2 of paddle. |
LARVA (fig. 6). Head: 0.95 mm. Siphon: 1.5-1.7 mm. Anal Saddle: 0.25 mm. As
figured, diagnostic characters as in the key. Readily separated from bisulcatus by
hair 6-I always triple; from arawak by the hairs 1-T and 2-II with more than 10
branches, always stellate. Head: Width about 1.1 of length. Hair 4-C single; 7-C with
9(8-11) and 9-C with 3 branches; 15-C short, double, not reaching base of mental
plate. Mental plate with a strong median tooth and 7 or 8 distinct teeth on either
side. Antenna: Length about 0.55 of head, with distinct spicules in basal 0.5; all
hairs single except 1-A (4-8). Thorax: Hair O-P usually with 18(18 or 19) stellate
branches; 7-P single; 14-P moderately long, single, narrow, attenuate apically; 1-M
with about 21 branches. Abdomen: Stellate hairs usually with at least 15 branches.
Hair 6-I triple; 2-II with at least 15 stellate branches; 2-I-VI distinctly cephalad of
hair 1. Segment VIII: Comb scales in a patch of 3 or 4 irregular rows, about 47-65 in
number; scales in proximal row fringed to apex, distal ending in a narrow spine. Si-
a Contrib. Amer. Ent. Inst., vol. 5, no. 1, 1969
phon: Index about 18.0-20.0. Integument moderately pigmented, faintly imbricate
- in basal 0.1. Subventral hairs moderately long, usually with less than 4 branches, pro-
gressively smaller caudad; subdorsal hairs long, single or double. Pecten extending to
0.35 of siphon; individual tooth long, with minute denticles restricted to ventral
side, almost extending to apex. Anal Segment: Integument moderately pigmented
and weakly spiculate; caudolateral marginal spines as in bisulcatus. Hair 1-X single;
2-X usually triple (3 or 4). Gills narrow, short, about 0.6 of dorsal saddle length.
SYSTEMATICS. C. antillummagnorum is undoubtedly a distinct species errone-
ously synonymized with bisulcatus (as americanus) by Rozeboom and Komp (1955:
77). Though similar to the closely related bisulcatus and arawak in adult coloration
and ornamentation, it is readily differentiated by constant characters of the male gen-
italia and immature stages. I have seen adults of antillummagnorum from Cuba, His-
paniola, Puerto Rico and the northern Virgin Islands (St. Thomas) and larvae from
Hispaniola, Puerto Rico and the northern Virgin Islands.
BIONOMICS. The immature stages of antillummagnorum have been found pri-
marily in the leaf axials of bromeliads and rarely in aroids at elevations of about 500
to 2,500 feet. Howard, Dyar and Knab (1915:306) state that W.V. Tower and J.H.
Pazos collected them (as Disulcatus) in artificial containers. These records have not
been confirmed and are probably erroneous. A few adults, including some blooded
females, have been collected by K.L. Knight in Cuba. According to Maldonado-Cap-
tiles, Pippin and Kuns (1958:58), adults of antillummagnorum (as americanus) rest
during the day in rock crevices and therefore presumably are either crepuscular or
nocturnal in activity. The statement of Pazos (Howard, Dyar and Knab, 1915:306)
that the adults (as bisulcatus) fly and bite during the day is probably due to a mis-
identification.
DISTRIBUTION (fig. 1). Known only from Cuba, Hispaniola, Puerto Rico and St.
Thomas in the northern Virgin Islands. Material examined: 182 specimens; 36 d, 55
2, 8 pupa, 83 larvae; 9 individual rearings (8 pupal, | larval).
CUBA. La Habana: San Antonio de los Banos, J.H. Pazos, 1 6 genitalia (786, slide no. 893)
[type no. 40778, USNM]. Locality not specified, 1914, JH. Pazos, 6 ? (782,783 ,787-789 865)
[USNM] . Oriente: Guantanamo Bay, adult collection, 7 July 1953, K.L. Knight, 2 2 (373); 27 Oct
1953, K.L. Knight, 7 6, 12 9 (451) [UCLA].
DOMINICAN REPUBLIC. Duerte: San Francisco, La Guama, elev. 1000 ft, 17 July 1968, D.C.
Watson (RDO 7) 3 L [UCLA]. La Vega: La Vega, elev. 300 ft, 25 July 1968, D.C. Watson (RDO
24),19,3P,1p,24L [UCLA].
HAITI. Trou Zombie, 26 May 1932, S.S. Cook, 1 6 with genitalia slide, 21 L [UCLA, USNM].
PUERTO RICO. Aguadilla: Maricao, 25 Aug 1935, G.S. Tulloch, 1 L (19209); 28 Jan 1936,
GS. Tulloch, 1 L (19332) [USNM]. Culebra Island: 21 Jan 1954, H.C. Hurt, 1 d, 3 9 (570) [UC-
LA]. Humacao: Dorado, Sardinera, 23 July 1932, 1 9 [USNM]. El Yunque, May 1942, 7d, 12°;
23 Jan 1943, H.D. Pratt, 7 L; 29 Sept 1943, H.D. Pratt, 7 L; 29 Sept 1943, H.D. Pratt and J.
Maldonado, 4 L; 25 June 1944, H.D. Pratt, 2 L [USNM]; elev. 500 ft, 11 Nov 1953, taro, K.L.
Knight, 3 6, 4 9 (524) [UCLA]. Luquillo Mts., elev. 2500 ft, 18 Feb 1935, J.G.N., 1 ¢ [USNM].
Mt. Briton, elev. 800-900 ft, 3 Aug 1964, bromeliads, K. and B. Bartholomew (PR 3), 1 L; 15 Aug
1964, K. and B. Bartholomew (PR 21), 1 Ip? (21-12), 1 L [UCLA] . Mayaguez: Mayaguez, 12 May
1936, colocasia axil, G.S. Tulloch, 1 6,3 L (19364); Sept (year not specified), W.V. Tower, 1 6,
1 3 genitalia (type no. 40778), 3 6, 5 2 [USNM]. Mona Island: 6 Oct 1954, W. Pippin, 1 d, 1?
[USNM]. San Juan: near San Juan, C.S. Ludlow, 1 2 [USNM]. Municipality not specified: El
Semil 31 May 1942, HD. Pratt, 1 6, 1 2 [USNM]. |
VIRGIN ISLANDS. St. Thomas: Charlotte Amalie, elev. 50-100 ft, 4 Mar 1969, M.M. Boreham
(VI 27), 3 pd (27-10,12,17), 5 p? (27-11,13-16), 2P, 1 p,6L, 51 [UCLA].
Berlin: Subgenus Micraedes of Culex | 37
3. Culex (Micraedes) arawak, Berlin, n.sp.
Figs. 1,7,10
TYPE: Holotype 6 (KO 29-19) with genitalia slide (680626-4), Moneague (St. Ann), Jamaica,
1943, R.B. Hill [USNM].
Culex (Aedinus) bisulcatus in part of Porter (1967:40); Stone (1967:218; 1969:7).
Culex (Aedinus) americanus in part of Stone, Knight and Starcke (1959:281).
Culex americanus of Hill and Hill (1945:2; 1948:52); Thompson (1947:78).
FEMALE. Unknown.
MALE. Wing: 3.0 mm. Proboscis: 1.8 mm. Forefemur: 2.2 mm. Similar to bisul-
catus differing only in the length of proboscis which is shorter than forefemur.
MALE GENITALIA (fig. 10). Diagnostic characters as in the key. Readily separ-
ated from other members of the group by the presence of only 6 or 7 apical teeth on
paraproct. Generally similar to bisulcatus differing only in the following. Ninth ter-
gite lobe broad, with 5 or 6 moderately long setae and without a clear distal portion.
Sidepiece length about 2.0 of greatest width; tergal surface with moderately long
bristles and 1 to several scattered scales; lateral and sternal surfaces without scales,
and with short bristles. Proximal division of subapical lobe prominent, with a long
narrow saber tergally and a flattened rod sternally; a smaller saber at base on tergal
surface; distal division with 3 or 4 relatively simple setae with hooked apex and |
longer seta with recurved apical barb. Clasper about 0.7 of sidepiece length, with a
long subbasal seta on external margin. Paraproct with only 6 or 7 apical teeth; cercal
setae 2.
PUPA. Unknown, but presumably similar to other members of group.
LARVA (fig. 7). Head: 0.8 mm. Siphon: 1.3-1.4 mm. Anal Saddle: 0.3 mm. As
figured; diagnostic characters as in the key. Readily separated from other members
of the group by the hair 2-II either single or double. Head: Width about 1.2 of
length. Hair 4-C single; 7-C with 5-9 and 9-C usually double (2 or 3); hair 15-C short,
double, not reaching base of mental plate. Mental plate with a strong median tooth
and 7 or 8 distinct teeth on each side. Antenna: Length about 0.5 of head, with spic-
ules in basal 0.5; all hairs single except 1(8-12). Thorax: Hair O-P usually with 16(14-
16) stellate branches; 7-P single; 14-P spikelike, single; 1-M with 10-14 branches. Ab-
domen: Stellate hairs usually with less than 15 branches. Hair 6-I with 4 branches; 2-
II spikelike, single or double; 2-I-VI slightly laterad of hair 1. Segment VIIT: Comb
scales in a patch of 3 irregular rows, 34-50 in number; all scales fringed to apex.
Siphon: Index about 10.5-12.0. Integument lightly pigmented, smooth. Subventral
hairs moderately long, at least 5 branched, progressively smaller caudad; proximal
subdorsal hair double, distal with 7 branches. Pecten extending to 0.35 of siphon; in-
dividual tooth long, with moderately long denticles on ventral side, extending almost
to apex. Anal Segment: Integument lightly pigmented, smooth; caudolateral border
with long spines as in bisulcatus. Hair 1-X single; 2-X usually triple (2 or 3). Gills
missing but presumably as in the other members of the group.
SYSTEMATICS. C. arawak has been confused in the past with bisulcatus (as amer-
icanus) but it is readily recognized by the characters in the diagnosis and in the keys.
Though similar to the members of the Bisulcatus Group in general characteristics,
arawak has the paraproct of the male genitalia with only 6 or 7 apical teeth, as in the
Schicki and Erethyzonfer groups. The association of larvae with the males is only
presumptive as no rearings have been made but it is probably correct since the im-
38 Contrib. Amer. Ent. Inst., vol. 5, no. 1, 1969
matures have been collected from the same general area as the males. This species ap-
pears to be a rare one for it has been found only twice in numerous collections made
in bromeliads in Jamaica in recent years.
BIONOMICS. The presumptive larvae of arawak have been collected in epiphytic
bromeliads at elevations of about 1,150 to 1,750 feet. Adults were found resting on
moist earth in limestone caves by Hill and Hill (1948:52).
DISTRIBUTION (fig. 1). Endemic to the island of Jamaica where it has been re-
ported only from the parish of St. Ann at an elevation of about 1,150 to 1,750 feet.
Material examined: 4 specimens; 2 6, 2 larvae; no individual rearings.
JAMAICA. St. Ann: cave near Moneague, 23 Jan 1943, R.B. Hill, 1 ¢ [UCLA]. Moneague,
1943, RB. Hill (KO 29-19), 1 6 [Holotype, USNM]; milestone 29 on Spanish Town Rd, elev.
1750 ft, 30 Aug 1967, W.A. Page, 1 L (JA 763); near Unity Valley, elev. about 1150 ft, 7 Sept
1967, W.A. Page, 1 L (JA 772) [UCLA].
SCHICKI GROUP |
FEMALES. Moderate in size. Head: Erect scales dark brown. Palpus short, about
0.2 of proboscis length; segment 4 about 3.0 length of 3. Thorax: Mesonotal scales
predominantly dark brown, appearing paler in caudal aspect; scales on anterior pro-
montory and humeral area white. Pleural integument mostly yellow; ppn, lower stp
and lower mep areas usually brown. Scales on upper margin of ppn white. Lower
mep area with 1 strong bristle. Abdomen: Sternites I-VI mainly creamy, with a few
apical dark scales on distal segments; sternite VII creamy basally and dark apically.
MALE (schicki only). Similar to female in coloration. Palpus long, about 0.9 of
proboscis; segment 3 long; segment 5 about 0.7 length of 4.
MALE GENITALIA (schicki only). Ninth tergite lobe moundlike, with 3-7 short
setae. Sidepiece roughly conical, length about 1.8-1.9 of greatest width; tergal sur-
face with 3 or 4 moderately long bristles at about middle; sternal surface with short-
er setae; no scales evident. Subapical lobe situated at 0.7, directed caudomesad;
proximal division with only 1 heavy rodlike appendage; distal division moderately
prominent, distinctly separate from proximal, bearing a prominent leaf in addition
to many relatively simple setae. Clasper about 0.6 of sidepiece length; distal 0.3 ex-
panded, curved inward, with a row of fine ridges on external margin and with 2 sub-
apical setae. Basal hook of lateral plate broadly sclerotized; distal part of apical pro-
cess rounded in tergal aspect; sternal spine projecting anteriorly. Paraproct with 6
apical teeth; cercal setae distinct, 8 or 9.
PUPAE. Cephalothorax: Hair 4-C short, usually single; 5-C moderately long but
shorter than or subequal to distance between its base and that of trumpet; 10-C usu-
ally with 3 or 4 branches. Abdomen: Hair 1-II with less than 10 branches, not re-
sembling float hair (1-I); 1-III-VIH varied in branching; 2-II-VI subequal to length of
9-II-VI; hair 2-VII distad of basal 0.80 of its tergite; 5-V,VI varied in length, sub-
equal to or about 1.5 length of succeeding segment; 9-III-V caudad of and 9-VI at
level of hair 6; hair 9-VII distinctly cephalad of 6, moderately long, either extending
barely or distinctly beyond apex of segment; 9-VIII moderately long, subequal in
length to segment VIII. Paddle: Elongate, slightly produced; margin with simple hair-
like spicules.
LARVAE. Head: Hair 4-C short to moderately long, at least double, distinctly
cephalolaterad of 5-C; hairs 8,10-C usually double; basal maxillary hair (bmh) doub-
le or triple, weak and attenuate apically. Antenna: Hair 1-A about 0.6 from base.
Thorax: Roughly rectangular in outline, wider than long. Hair 4-P double; 9-M at
Berlin: Subgenus Micraedes of Culex 39
least 4 branched (4-8). Abdomen: Hairs 1,2,4,11,13-I, 1,5,9,13-H-VI, 5,8,13-VII with
a few to many simple branches; 2-II-VI weakly stellate; 2-I,II cephalolaterad and 2-
III-VI cephalomesad of hair 1. Segment VIII: Individual scale fringed to apex, with-
out a median spine. Hairs 1,5-VIII with a few to many simple branches. Siphon:
Long, index about 9.0-12.0. Pecten terminating before middle. Subventral hairs long,
progressively smaller distally; proximal subdorsal hair distad of hair 1-S and usually
caudad of last pecten tooth. Anal Segment: Caudolateral border of saddle with mod-
erately long marginal spines, no distinct spines ventrad of hair 1-X. Hair 1-X short,
forked, at least 4 branched. Saddle body imbricate, without spicules. Hair 2-X doub-
le (with 4-6 branches in sandrae). Gills long, subequal, at least 2.0 of dorsal saddle
length.
DISCUSSION. The Schicki Group is recognized here for the nominate form and
sandrae. The recognition of this group is primarily based on the nature of the male
genitalia, although differences in the adults and the immature stages also warrant
their separation from the Erethyzonfer Group. The adults are separated from other
members of the subgenus by the length of palpus and the presence of white scales on
the upper margin of ppn. The larvae are diagnosed by the absence of stellate hairs on
the body and by the forked hair 1-X of the saddle. The pupae are readily separated
from the Bisulcatus Group by the presence of marginal spicules on the paddle, a fea-
ture which they share with the Erethyzonfer Group.
The Schicki Group is known at present only from the Sierra Madre del Sur in
southern Mexico (fig. 1), where both breed in epiphytic bromeliads, schicki at eleva-
tions of about 2,200 feet and sandrae at elevations of 7,000 to 8,000 feet.
4. Culex (Micraedes) schicki Berlin, n.sp.
Figs. 1,8,9
TYPES: Holotype 6 (MEX 415-11), with associated pupal and larval skins and genitalia slide,
Rancho Viejo de Agua de Obispo, Chilpancingo, Guerrero, Mexico, elevation about 2,200 ft, larva
from epiphytic bromeliads, 8 Aug 1966, D.A. Schroeder [USNM]. Allotype 2? (MEX 417-107),
with associated pupal skin, same data as holotype [USNM] . Paratypes: 1 Ipd (415-10), 1 lp? (415-
12), 1 L (416), 1 lpd (417-27), 1 lp? (417-40), 1 lp (417-41), 2 P,5 L, 11(417), 1 L (418), 1 Ip?
(421-13), 1 p? (421-100), 7 L (421), 1 lpd (427-12), 1 lp? (427-11), 1 pd (427-101), 1 IP (427-
13), same data as holotype [USNM, UCLA]. This species is dedicated to Robert X. Schick in rec-
ognition of his contributions to the knowledge of Neotropical mosquitoes.
FEMALE. Wing: 3.4 mm. Proboscis: 2.1 mm. Forefemur: 1.9 mm. Abdomen:
2.4-2.5 mm. As described for the subgenus and the group and with the following di-
agnostic features. Decumbent scales on dorsum of head whitish. Proboscis longer
than forefemur. Mesonotal scales auburn, linear. Scales on antealar area above para-
tergite mainly auburn, some whitish anteriorly. Basolateral light patches on abdom-
inal tergites whitish.
MALE. Wing: 2.9 mm. Proboscis: 1.95 mm. Forefemur: 1.6 mm. Similar to fe-
male in coloration. Proboscis longer than forefemur.
MALE GENITALIA (fig. 8). Diagnostic characters as in the key. Readily separat-
ed from other members of the subgenus by the presence of only | rodlike appendage
on the proximal division of the subapical lobe. Ninth tergite lobe prominent, with 3-
7 weak setae. Sidepiece length about 1.9-2.0 of greatest width; a group of 3 or 4
moderately long bristles tergolaterally and a few long bristles laterally; other setae
40 Contrib. Amer. Ent. Inst., vol. 5, no. 1, 1969
on tergal surface weaker, confined to middle and distolateral surfaces. Proximal divi-
sion of subapical lobe prominent, split apically into 2 unequal parts, longer bearing a
broad saber with recurved apex, shorter a small saber, 2 small setae arising from base
of tergal surface of basal part; distal division with at least 16 short to moderately
long, relatively simple setae, most of them with recurved apices, and with a promin-
ent leaf and a narrow seta on base of tergal surface. Clasper about 0.65 of sidepiece
length; distal 0.3 bent inward. Paraproct with 6 apical teeth; cercal setae 8 or 9.
PUPA (fig. 8). Abdomen: 2.5 mm. Trumpet: 0.45 mm; index about 9.0. Paddle:
0.7 mm. As figured; diagnostic characters as in the key. Readily separated from sand-
rae by very long hair 6-II, at least 2.0 of 7-II. Cephalothorax: Lightly to moderately
pigmented, lateral areas darker. Hair 1-C usually double (1-3); hair 5-C triple (3-5);
hairs 6,7,9-C single; 8-C double (2-3); hair 11-C usually single (1-4); hair 12-C triple
(2-4). Abdomen: Integument moderately pigmented, progressively lighter caudad.
Hair 1-I with 6-11 branches; 6-I,II single, long, at least 2.0 of 7-I,II; hair 5-V,VI usu-
ally double or triple, at least 1.5 of succeeding segment; 6-IV,V long, extending to
apex of succeeding segment; 6-VII usually triple (2-4), short, about 0.5 of 9-VII;
hair 9-VIII usually triple (2-3). Paddle: Elongate, longer than wide, at least 2.5 of
segment VIII. Male genital lobe extending to 0.43 and female to 0.2 of paddle.
LARVA (fig. 9). Head: 0.85 mm. Siphon: 1.4 mm. Anal Saddle: 0.3 mm. As fig-
ured; diagnostic characters as in the key. Readily separated from sandrae by the
double or triple hair 2-X and the short proximal subdorsal hair of the siphon (about
0.25 of 1-S in length). Head: Width about 1.05-1.1 of length. Hair 4-C usually 4
branched (3-6); hair 7-C with 5-10 branches; 9-C usually 11 branched (9-16); hair
15-C usually double, moderately long, barely extending beyond base of mental plate.
Mental plate with a strong median tooth and 7 or 8 distinct teeth on either side. An-
tenna: Length about 0.5 of head, with distinct spicules in basal 0.6; all hairs single
except 1-A (3-9). Thorax: Hair O-P with 11-18 branches; 7-P always double; 14-P
single; 1-M usually with 12(12-21) branches. Abdomen: Hair 2-II-VII multiple, weak-
ly stellate, with at least 5(5-16) branches; 6-I usually double (2-4); hair 9-I-VI at least
4 branched. Segment VIII: Comb scales in a patch of 3 irregular rows, about 34-37
in number. Siphon: Index about 9.0. Integument moderately pigmented, darker api-
cally; imbrication distinct in basal 0.3. Distal subventral hair small, subequal to sub-
dorsals; other subventral hairs long, strongly pigmented; proximal subdorsal hair
small, double or triple, about 0.25 length of 1-S. Pecten extending to basal 0.3; indi-
vidual tooth short, with a weak subbasal ventral tooth. Anal Segment: Integument
of saddle imbricate, moderately to strongly pigmented; caudolateral border with
moderately long spines. Hair 1-X short, forked; 2-X usually double, rarely triple.
Gills moderately broad, 2.5-3.5 of dorsal saddle length.
SYSTEMATICS. The adults of schicki cannot be differentiated from sandrae, but
the immature stages are strikingly distinct, as indicated in the diagnosis and in the
keys. !
BIONOMICS. The immature stages of schicki have been collected in epiphytic
bromeliads. All known adults are reared and nothing is known about the behavior of
the adults.
DISTRIBUTION (fig. 1). Province of Gerrero in southern Mexico at elevations of
about 2,200 feet. Material examined: 51 specimens; 5 6, 6 9, 15 pupae, 25 larvae; 13
individual rearings (3 pupal, 8 larval, 2 incomplete).
MEXICO. Guerrero: Chilpancingo, Rancho Viego de Agua de Obispo, elev. 2200 ft, 8 Aug
1966, D.A. Schroeder (MEX 415-418,421,427, type series), 2 lpd (415-10,11), 1 Ip? (415-12), 1L
(416), 1 Ipod (417-27), 1 lp? (417-40), 1 p? (417-107), 1 Ip (417-41), 2 P, 5 L, 11(417), 1
Berlin: Subgenus Micraedes of Culex 41
L (418), 1 lp? (421-13), 1 p? (421-100), 7 L (421), 1 Ipd (427-12), 1 Ip? (427-11), 1 pé (427-
101), 1 IP (427-13) [USNM, UCLA].
5. Culex (Micraedes) sandrae Berlin, n.sp.
Figs. 1,10,11
TYPES: Holotype ° (MEX 362-100), with associated pupal skin, Omilteme, Guerrero, Mexico,
elevation 8,000 ft, pupa from epiphytic bromeliad, 8 Sept 1965, D.A. Schroeder [USNM]. Para-
types: 1 IP (362-10), 18 L (362), same data as holotype [USNM, UCLA] . This species is dedicated
to Miss Sandra J. Heinemann in recognition cf her contributions to the knowledge of Neotropical
mosquitoes. ;
FEMALE. Wing: 3.4 mm. Proboscis: 2.05 mm. Forefemur: 1.7 mm. Abdomen:
2.3 mm. Essentially similar to schicki from which it cannot be separated.
MALE. Unknown.
PUPA (fig. 10). Abdomen: 2.8 mm. Trumpet: 0.5 mm; index about 9.0. Paddle:
0.75 mm. As figured; diagnostic characters as in the key. Description based on a
single skin. Readily separated from schicki by the moderately long hair 6-II which is
subequal to or slightly longer than 7-II. Cephalothorax: Lightly to moderately pig-
mented, lateral areas darker. Hair 1-C single; 5-C triple; 6-9-C usually single; 11-C
double; 12-C triple. Abdomen: Integument moderately pigmented, progressively
lighter caudad. Hair 1-II with 4 branches; 6-I,II single, moderately long, subequal to
or slightly longer than 7-I,IJ; hair 5-V,VI double or triple, barely extending to apex
of succeeding segment; 6-III-VI moderately long, extending to 0.5 of succeeding seg-
ment; 6-VII double, short, about 0.5 of 9-VII; hair 9-VIII missing, probably triple.
Paddle: Elongate, longer than wide, at least 2.0 of segment VIII. Female genital lobe
extending to 0.2 of paddle.
LARVA (fig. 11). Head: 0.9 mm. Siphon: 1.65 mm. Anal Saddle: 0.33 mm. As
figured; diagnostic characters as in the key. Readily separated from schicki by the
more numerous branches (4-6) in hair 2-X and the long proximal subdorsal hair of
the siphon (subequal in length to hair 1-S). Head: Width about 1.05-1.1 of length.
Hair 4-C double or triple; 7-C usually with 7(7-11) branches; 9-C usually with 7
branches (6-12); hair 15-C short, double, terminating before base of mental plate.
Mental plate with a strong median tooth and 7 distinct teeth on either side. Anten-
na: Length about 0.5 of head, with distinct spicules in basal 0.6; all hairs single ex-
cept 1(6, 4-7). Thorax: Hair O-P with 7-11 branches; 7-P single; 14-P single; 1-M usu-
ally with 7(6-8) branches. Abdomen: Hair 2-I-VII simple, usually triple; 6-I at least
triple (3-5); hair 9-I-VI single or double. Segment VIII: Comb scales in a patch of 3
irregular rows, about 36-50 in number. Siphon: Index about 10.0-12.0. Integument
moderately pigmented, slightly darker apically; imbrication distinct in basal 0.3.
Distal subventral hair small, subequal to distal subdorsal hair; other subventral hairs
long, strongly pigmented; proximal subdorsal hair long, 3 or 4 branched, usually sub-
equal to 1-S. Pecten extending to basal 0.3; individual tooth as in schicki. Anal Seg-
ment: Integument of saddle imbricate, moderately to strongly pigmented; caudolat-
eral border with spines as in schicki. Hair 1-X short, forked; 2-X with at least 4(4-
6) branches. Gills moderately broad, 2.4-2.8 of dorsal saddle length.
SYSTEMATICS. C. sandrae is recognized here as a distinct species entirely on the
basis of striking constant features in the immature stages. The association of the
known stages is probably correct although no complete larval rearings were made.
42 ) Contrib. Amer. Ent. Inst., vol. 5, no. 1, 1969
BIONOMICS. The immature stages of sandrae have been collected in the leaf axils
of epiphytic bromeliads. Nothing is known about the behavior of adults.
DISTRIBUTION (fig. 1). Provinces of Guerrero and Oaxaca in southern Mexico at
elevations of about 7,200-8,000 feet. Material examined: 41 specimens; 1 ?, 2 pu-
pae, 38 larvae; 2 individual rearings (1 pupal, 1 incomplete).
MEXICO. Guerrero: Omilteme, elev. 8000 ft, 8 Sept 1965, D.A. Schroeder (MEX 362, type ser-
ies), 1 p? (362-100), 1 IP (362- 10), 18 L (362) [USNM, UCLA]. Oaxaca: Ixtlan de Juarez, Vivero
Rancho Teja, elev. 7200 ft, 12 Aug 1966, D.A. Schroeder, 4 L (MEX 429), 15 L (MEX 430)
[UCLA].
ERETHYZONFER GROUP
FEMALE. Moderate in size. Head: Erect scales dark brown to black. Palpus short,
about 0.25-0.28 of proboscis length; segment 4 about 3.0 length of 3. Thorax: Mes-
onotal scales predominantly brown; scales on anterior promontory and humeral area
white. Pleural integument yellowish. Scales on upper margin of ppn brown. Lower
mep area with 1 strong bristle. Abdomen: Sternites I-VI predominantly creamy, with
a few apical dark scales on distal segments; sternite VII creamy basally and dark api-
cally.
MALE. Similar to female in coloration. Palpus long, subequal to proboscis in
length and 5-segmented; segment 5 about 0.7-0.75 length of 4.
MALE GENITALIA. Ninth tergite lobe distinct, moundlike, with at least 12 mod-
erately long setae. Sidepiece roughly conical, length about 1.7-1.8 of greatest width;
with a large patch of long bristles tergally; sternal surface with short to moderately
long setae; no scales evident. Subapical lobe in distal 0.6, directed mesad; proximal
division with 2 heavy subequal rods; distal division sessile, separate from proximal,
bearing in addition to relatively simple setae, | leaf and 1 lanceolate seta on tergal
surface. Clasper about 0.67 of sidepiece length; distal 0.3 expanded, curved inward,
with a row of fine ridges on external margin and with 2 subapical setae. Basal hook
of lateral plate broadly sclerotized; distal part of apical process rounded in tergal as-
pect; sternal spine projecting anteriorly. Paraproct with 6 or 7 apical teeth; cercal
setae distinct, 2 or 3.
PUPA. Cephalothorax: Hair 4-C short, usually double; 5-C moderately long, sub-
equal in length to distance between its base and that of trumpet; hair 10-C usually
with 7(3-14) branches. Abdomen: Hair 1-II with at least 10 branches, sometimes
faintly resembling float hair (1-1); hair 1-III-VII varied in branching; 2-II-VI subequal
in length to 9-II-VI; hair 2-VII distad of basal 0.80 of its tergite; 5-V,VI long, ex-
tending beyond apex of succeeding segment; 9-II-V caudolaterad and 9-VI at level of
hair 6; hair 9-VII distinctly cephalad of 6, extending beyond apex of segment; 9-VIII
moderately long, subequal in length to segment VIII. Paddle: Elongate, slightly pro-
duced; margin with simple, hairlike spicules.
LARVA. Head: Hair 4-C short to moderately long, usually double, slightly cephal-
olaterad of 5-C; hairs 8,10-C double; basal maxillary hair (bmh) small, single and at-
tenuate apically. Antenna: Hair 1-A about 0.5 from base. Thorax: Roughly oval in
outline, wider than long. Hair 4-P usually double, rarely triple; 9-M with 5 or 6
branches. Abdomen: Hairs 1,2,4,11,13-I, 1,2,5,9,13-II, 1,2,5,7,9,13-III-VI, 1,2,5,8,
10,13-VII distinctly stellate, with many spikelike branches. Hair 2-I distinctly cep-
halolaterad of hair 1; hair 2-II-V cephalomesad and 2-VI cephalad of hair 1. Segment
VII: Individual scale long, fringed on sides and terminating in a long spine. Hair 1-
VIII stellate. Siphon: Long, index about 14.0-16.0. Pecten extending to 0.3. Subven-
Berlin: Subgenus Micraedes of Culex 43
tral hairs long, progressively smaller distally; proximal subdorsal hair distad of hair 1|-
S and usually caudad of last pecten tooth. Anal Segment: Caudolateral border of
saddle with long marginal spines, some with lateral spinules; a few distinct spines
ventrad of hair 1-X. Hair 1-X moderately long, usually double. Saddle body with
rows of small spicules. Hair 2-X double. Gills long, subequal, about 2.0 of dorsal sad-
dle length.
DISCUSSION. The Erethyzonfer Group includes only the nominate form. The
adults differ from other members of the subgenus in the length of the palpus in the
2 sexes, in the female about 0.25-0.28 and in the male subequal to the proboscis
length. The male genitalia are diagnostic by having 1 leaf and 1 lanceolate seta on
the subapical lobe of the sidepiece in addition to other relatively simple setae. The
larvae are separated from the other groups by hair 1-X being double and by the pres-
ence of distinct spines ventrad of hair 1-X on saddle.
The Erethyzonfer Group is Central American in distribution. To date it has been
collected only in Guatemala, Costa Rica and Panama but it probably occurs in suit-
able environments in intermediate areas. The immature stages have been collected
primarily in the leaf axils of epiphytic and terrestrial bromeliads and rarely in aroids.
6. Culex (Micraedes) erethyzonfer Galindo & Blanton
Figs. 1,12,13
1954. Culex (Microculex) erethyzonfer Galindo and Blanton, 1954:244-246. TYPE: Holotype ¢
with genitalia slide, Palo Santo, Chiriqui Volcano region (Chiriqui), Panama, elevation
4,500 ft, 18 Oct 1950, P. Galindo [USNM].
Culex (Microculex) erethyzonfer of Stone and Knight (1957:58); Stone, Knight and Starcke
(1959:279); Belkin, Schick and Heinemann (1965:55).
FEMALE. Wing: 3.5 mm. Proboscis: 2.1 mm. Forefemur: 1.9 mm. Abdomen: 2.5
mm. As described for the subgenus and the group and with the following additional
features. Decumbent scales on dorsum of head narrow, whitish. Proboscis longer than
forefemur. Mesonotal scales dark brown, linear. Antealar area above paratergite with
scattered dark brown scales. Basolateral light patches on abdominal tergites creamy.
MALE. Wing: 3.2 mm. Proboscis: 2.1 mm. Forefemur: 1.8 mm. Similar to female
in general features.
MALE GENITALIA (fig. 12). Diagnostic characters as in the key. Readily separ-
ated from other members of the subgenus by the prominent ninth tergite lobe bear-
ing at least 12 moderately long setae and by the distal division of the subapical lobe
being sessile and bearing | prominent leaf and 1 lanceolate seta on the tergal surface
in addition to other setae. Sidepiece length about 1.7-1.8 of greatest width; a patch
of long bristles tergally and longer bristles laterally; sternal surface with smaller set-
ae. Proximal division of subapical lobe prominent, split apically into 2 equal parts,
each bearing a rod, larger rod with a recurved apex, 3 small simple setae arising from
base of tergal surface; distal division with 18-22 short to moderately long, relatively
simple setae, most of them with recurved apices, and with a prominent leaf and a
distinctly lanceolate seta on base of tergal surface. Clasper about 0.67 of sidepiece
length; distal 0.3 curved inward. Paraproct with 6 or 7 apical spines; cercal seta 2.
PUPA (fig. 12). Abdomen: 2.6 mm. Trumpet: 0.5-0.55 mm; index about 11.0-
12.0. Paddle: 0.7 mm. As figured; diagnostic characters as in the key. Readily separ-
44 Contrib. Amer. Ent. Inst., vol. 5, no. 1, 1969
ated from members of the Bisulcatus Group by the presence of marginal spicules on
the paddle; from schicki by the shorter hair 6-IV,V and 9-VIII usually 4 branched;
from sandrae by hair 1-I with at least 10 branches. Cephalothorax: Integument light-
ly pigmented. Hair 1-C single; 5-C usually triple (2-5); hairs 6,7 9-C single; 8-C usual-
ly double (2-4); hair 11-C single; 12-C usually with 4(3-6) branches. Abdomen: Inte-
gument lightly pigmented. Hair 1-I with at least 10 branches (10-25); hair 6-I,]] mod-
erately long, about 1.5 of 7-I,II; hair 5-V,VI usually 2-4 branched, about 1.5 of suc-
ceeding segment, rarely subequal to segment length; 6-IH-VI moderately long, ex-
tending to 0.5 of succeeding segment; 6-VII usually. triple (2-5), about 0.4 of 9-VIT;
hair 9-VIII usually with 4(2-5) branches. Paddle: Oval, longer than wide, about 2.5
of segment VIII. Male genital lobe extending to 0.43 and female to 0.2 of paddle.
LARVA (fig. 13). Head: 0.8 mm. Siphon: 1.5 mm. Anal Saddle: 0.25 mm. As fig-
ured; diagnostic characters as in the key. Readily separated from other members of
the subgenus by the presence of distinct marginal spines ventrad of hair 1-X on sad-
dle. Head: Width about 1.2 of length. Hair 4-C usually double (1-4); hair 7-C with 5-
10 and 9-C with 8-14 branches; 15-C double (2-4), barely extending to base of men-
tal plate. Mental plate with a strong median tooth and 7 or 8 distinct teeth on either
side. Antenna: Length about 0.6 of head, with distinct spicules in basal 0.6; all hairs
single except 1(6, 5-9). Thorax: Hair O-P with at least 15 branches (15-23); hair 7-P
single; 14-P single; 1-M with 17-39 branches. Abdomen: Hair 2-I-VII with at least 9
branches (9-29); hair 6-I usually triple (2-4); hair 9-I usually triple (2-5); hair 9-II-VI
with at least 5(5-10) spikelike branches. Segment VIII: Comb scales in a patch of 3
or 4 irregular rows, 32-52 in number. Siphon: Index about 14.0-16.0. Integument
lightly to moderately pigmented, faintly imbricate in basal 0.2. Distal subventral
hair short, subequal to distal subdorsal hair; other subventral hairs long, moderately
pigmented; proximal subdorsal hair long, subequal to hair 1-S. Pecten usually ex-
tending to 0.3 and terminating before hair 1-S, in some specimens, | or 2 teeth dis-
tad of 1-S; individual tooth with a few small denticles, restricted to ventral side. Anal
Segment: Integument of saddle lightly to moderately pigmented, uniformly spicu-
late; caudolateral margin with long spines extending ventrad of hair 1-X. Hair 1-X
moderately long, usually double (1-3); hair 2-X usually double (1-2). Gills narrow,
about 2.0 of dorsal saddle length. |
SYSTEMATICS. C. erethyzonfer is easily differentiated in all stages from all other
members of the subgenus by the characters in the diagnosis and in the keys. The des-
cription of erethyzonfer is based on associated topotypic material from Panama.
Very little variation is apparent in the large series of specimens examined except that
some larvae from Panama, Costa Rica and Guatemala have 1 or 2 pecten teeth dis-
tinctly beyond hair 1-S on the siphon.
BIONOMICS. The immature stages of erethyzonfer have been reported from the
leaf axils of epiphytic and terrestrial bromeliads and rarely from aroids at elevations
above 4,500 feet. The holotype was taken in a sweeping collection.
DISTRIBUTION (fig. 1). Highlands of Guatemala (probably Honduras, El Salva-
dor and Nicaragua), Costa Rica and Panama. Material examined: 486 specimens; 33
3, 46 2, 105 pupae, 302 larvae; 89 individual rearings (18 pupal, 45 larval, 26 incom-
plete).
COSTA RICA. Alajuela: Ciruelas, 13 Nov 1962, ?ground pool, C.L. Hogue and W.A. Powder,
(CR 16), 1 6 [UCLA]. Cartago: near Cervantes, 9 Nov 1962, bromeliads, C.L. Hogue (CR 12,13),
5 Ipd (12-103,109,116,119,120), 6 Ip? (12-104,108,111,112,114,118), 3 pd (12-102,107,117),
5 p? (12-101,113,121-123), 1 IP (12-115), 1 9, 1 P, 1 p, 14. L, 71 (12), 1 Ipé (13-208), 6 Ip? (13-
107,201-204,207), 3 L (13) [UCLA]. La Sierra, elev. 6300 ft, 24 Nov 1962, bromeliads, C.L.
Hogue and W.A. Powder (CR 54), 1 Ipd (54-108), 1 pd (54-201), 3 6, 3 9, 11 P, 19 L [UCLA].
Berlin: Subgenus Micraedes of Culex 45
GUATEMALA. Alta Vera Cruz: Trece Aguas, 7 July 1964, J.E. Zavortink (GUA 18), 2 L; 7
July 1964, J.E. and T.J. Zavortink (GUA 19), 1 lpd (19-21), 1 Ip? (19-25), 2 L [UCLA].
PANAMA. Bocas del Toro: La Sierra, elev. 5400 ft, 8 Apr 1963, bromeliads, A. Quinonez (PA
180), 1 P, 31 L; 8 Apr 1964, shannon trap, A. Quinonez (PA 182), 1 9 [UCLA]. La Zorra, elev.
4400 ft, 5 Apr 1963, bromeliads, A. Quinonez (PA 172,173), 2 IP (172-101,103), 1 Ip? (173-102),
1 pd (173-108), 1 IP (173-116), 5 L (173); 6 Apr 1963, bromeliads, A. Quinonez (PA 176,177),
4 L (176), 2 L (177); 6 Apr 1963, leaf axil of aroid, A. Quinonez (PA 178), 1 L; 7 Apr 1963,
bromeliads, A. Quinonez (PA 179), 9 L [UCLA]. Chiriqui: Bambito, elev. 5500 ft, 20 Oct 1945,
P. Galindo, 1 pd (00103), 5 L [paratypes, USNM]. Cerra Punta, elev. 5000 ft, 26 May 1946, P.
Galindo, 1 ¢ (00102), 1 6 genitalia (00104) [paratypes, USNM]. El Hato, elev. 4000 ft, 5 Sept
1946, P. Galindo, 1 6 genitalia (00101) [paratype, USNM]. Hato del Volcan, Chiriqui Lagoon,
elev. 4500 ft, 12 Mar 1964, bromeliads, A. Quinonez (PA 638), 9 L [UCLA]. Darien: Pucro,
Cerro Mali, elev. 4800-5000 ft, 22 May 1963, bromeliads, A. Quinonez (PA 348), 3 lp? (348-103,
105,109), 2 IP (348-108,112), 2 L; 23 May 1963, bromeliads, A. Quinonez (PA 357), 1 Ipé (357-
109), 1 Ip? (357-111), 1 IP (357-108), 1 P, 1 L; 24 May 1963, bromeliads, A. Quinonez (PA 359),
3 Ipd (359-138,140,160), 6 lp? (359-142,144,146,150,158,159), 1 1d (359-151), 2 pd (359-104,
111), 2 p? (359-147,153), 5 IP (359-145 ,148,152,156,157), 69 L, 1 1;26 May 1963, bromeliads,
A. Quinonez (PA 363), 2 Ipd (363-107,131), 5 lp? (363-108,109,114,134,135), 1 pd (363-113),
1 p? (363-112), 4 IP (363-120,133,136,137), 33 L, 11; 1 June 1963, bromeliads, A. Quinonez
(PA 371), 3 Ip? (371-113,120,126), 1 19 (371-111), 4 Ip (371-102,107,112,129), 2 IP (371-116,
131), 5 L, 1 1; 3 June 1963, bromeliads, A. Quinonez (PA 373), 2 Ipé (373-107,111), 3 IP (373-
103,109,110); 7 June 1963, wooden bowl, A. Quinonez (PA 377), 1 L [UCLA].
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O COND NN BR WN
Berlin: Subgenus Micraedes of Culex 49
FIGURES
Distribution of groups and species of Micraedes
Culex (Micraedes) bisulcatus; adult morphology
Culex (Micraedes) bisulcatus; male genitalia and pupa
Culex (Micraedes) bisulcatus; larva
Culex (Micraedes) antillamimagnorum: male conliale and pupa
Culex (Micraedes) antillummagnorum; larva
Culex (Micraedes) arawak:; larva
Culex (Micraedes) schicki; male genitalia and pupa
Culex (Micraedes) schicki; larva
Culex (Micraedes) sandrae ; pupa. Culex (Micraedes) arawak; male genitalia
Culex (Micraedes) sandrae; larva
Culex (Micraedes) erethyzonfer; male genitalia and pupa.
Culex (Micraedes) erethyzonfer; larva
HIN
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Fig. 3
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Route de Traversee
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MICRAEDES
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MICRAEDES _ ;
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MICRAEDES
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MICRAEDES
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MICRAEDES
Fig. lO
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Guerrero
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sidepiece
MICRAEDES
sandrae
MEX 362
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Guerrero
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Cs
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MICRAEDES
Fig. I2
subapical lobe
?
{ erethyzonfer \,
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ed
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Cervantes ly
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Costa Rica
3
MICRAEDES
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Berlin: Subgenus Micraedes of Culex
INDEX TO SCIENTIFIC NAMES
he
Aedes (Howardina), 23, 24, 30
Aedinus, 24
Aedinus of authors, 22, 24
americanus of authors, 22, 30, 31, 32, 35, 36, 37
Anoedioporpa, 24
Anopheles (Kerteszia), 23
antillummagnorum, 22, 23, 24, 26, 27-29k, 30, 31, 32, 34-36; /, 5, 6
arawak, 23, 24, 25, 27-29k, 29, 30, 31, 35, 37-38; J, 7, 10
bisulcatus, 21, 22, 24, 26, 27-29k, 30, 30-34, 35, 36, 37; /, 2, 3,4
bisulcatus of authors, 35, 37
Bisulcatus Group, 24, 25, 26, 27-29k, 29-30, 37, 39, 44; /
Culex, 22, 23, 24
Culex (Microculex), 23
erethyzonfer, 27-29k, 43-45; /, 12, 13
Erethyzonfer Group, 24, 27-29k, 37, 39, 42-43; /
~ Eubonnea, 24
Melanoconion, 22, 24
Micraedes, 21, 22, 23, 24-27; /
Microculex, 24
sandrae, 24, 27-29k, 39, 40, 41-42, 44; 7, 10, 11
schicki, 24, 27-29k, 38, 39-41, 41, 44;/,5,9
Schicki Group, 24, 27-29k, 37, 38; J
Tinolestes, 22, 24
Toxorhynchites (Lynchiella), 23
Wyeomyia, 23
63
ay
her |
a
i
roe
A,
3
=x
=
ae
(Continued from inside front cover)
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MOSQUITO STUDIES (Diptera, Culicidae)
XIX. The treehole Anopheles of the New World
By Thomas J. Zavortink
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(Continued on back cover)
MOSQUITO STUDIES (Diptera, Culicidae)
XIX. THE TREEHOLE ANOPHELES OF THE
NEW WORLD’
By
Thomas J. Zavortink’
CONTENTS
EGS Fa 9 (08 aR BRP IG Pt Rar aR a ee ee Ser mn nae an RC RH
Bo ER EICS iiss ayn, ss Pedaimiatars iia ONE hl mkt tk are tilde hileie tihagiisias v
TAXONOMIC TREATMENT |
Deepen iat al Clean we) wiih biranr rh UN tee gle ly atl ol atl le a ol)
Kevsdo- Species). ia. OG RR es SIR ME EET eee
1. Anopheles (An.) havent Conuilictt . Ca WHE he TG: Mas tL, alae Loo
pul wowneies (Amt Iuditine Zaviir Osis tee ig vase moe A Os oak a Gla. PLO
cE CLee I AU al AUST WAPOA i i cu ken aia aisha en mate vasa, RS
Mc AHOUUCIEG. A LOFOGUICGIUA: DISD. (ot 6 i) Bed hk keke Bile ae) eo a Le
Somnopreies (An bxolmiuensis de UeONss cee 6G ar Rew eb ee AD
6. Anopheles (An. be ale i aie Pry iiie M aorahida) «Mak ua hoah Bali oh Sakae
Pere neNChs CUED... oS UR LR et hee Aah ad Re Rta Ig, fy
See eye rr Lda iti! Ca Math en fae Same uan ire Uw slice br ade
PAB AD 4.) Piha ae Posie Cray, Gone Br umUAaayy ay beck ae
INDEX TO SCIENTIF IC NAMES Se GM i te ed fa ETO ig ae oe Re a es
INTRODUCTION
The present paper is a review of the obligate treehole breeding Anopheles of the
New World. It is essentially an expansion of my previous study of the treehole
Anopheles of the United States (Zavortink, 1969).
The taxonomic procedure is that outlined for the project “Mosquitoes of Middle
America” by Belkin, Schick et al (1965:10-11). Format, abbreviations and other
details follow earlier papers in this series. I have not seen the types of barberi, fausti,
or xelajuensis.
I am indebted to the following individuals for loans of specimens included in this
‘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.
ps | Contrib. Amer. Ent. Inst., vol. 5, no. 2, 1969
study: Alfonso Diaz Najera of the Instituto de Salubridad y Enfermedades Tropi-
cales, Mexico [ISET]; Lewis T. Nielsen of the University of Utah [UTAH]; L.L.
Pechuman of Cornell University [CU]; Lloyd E. Rozeboom of Johns Hopkins Uni-
versity [HOPK]; and Alan Stone of the United States National Museum [USNM].
Peter F. Mattingly kindly loaned specimens of extralimital species from the British
Museum (Natural History) collection.
I thank John N. Belkin for editing the manuscript and the following individuals,
all present or past members of the “‘Mosquitoes of Middle America” project staff,
for technical assistance: Caryle Abrams, Sheila Bernstein, Sally Dieckmann, Frances
Dupont, Sandra Heinemann, Christopher Ishida, L. Margaret Kowalczyk, Nancy
Martsch, Michael Nelson and William Powder.
SYSTEMATICS
The American treehole breeding Anopheles, barberi Coquillett, 1903, fausti Var-
gas, 1943, judithae Zavortink, 1969, xelajuensis de Leon, 1938, and 2 species des-
cribed here, arboricolus and powderi, belong to a single phylad. At least 2 of the
numerous Old World species of container breeding Anopheles, barianensis James,
1911 and plumbeus Stephens, 1828, belong to this same line. This group of 8
species is differentiated morphologically and biologically, by its usual restriction to
_ temperate or montane tropical regions, from the other phylads of container breed-
ing anophelines.
As members of this group have some rather unusual features in the preimaginal
stages, 3 generic group taxa have been proposed for included species: Coelodiazesis
Dyar & Knab, 1906 founded on larval characteristics of barberi; Cyclophorus Eysell,
1912 on the immature stages of plumbeus; and Russellia Vargas, 1943 on larval pe-
culiarities of xelajuensis. Despite its distinctiveness, this group of species is not cur-
rently recognized at the subgeneric level and is placed in Anopheles (Anopheles)
(Stone, Knight and Starcke, 1959). This is because anophelines of several distinct
phylads adapted to breeding in plant containers show convergence in some of the
more conspicuous elements of the larval chaetotaxy, such as reduction of the front-
al hairs of the head and elongation of lateral abdominal hair 6-IV-VI, and, as a con-
sequence, students of the genus have been reluctant to use features of the immature
stages to define or place them.
On the basis of morphology of the adult, male genitalia and larva, geographical
distribution and habitat, the 6 species included here fall into at least 3 different
primary groups: 1) barberi and judithae, 2) fausti and arboricolus and 3) xelajuensis.
The affinities of the poorly known powderi are uncertain. The geminate species bar-
beri and judithae differ most conspicuously from the others in the following fea-
tures: as adults by 1) interocular scales absent or short, 2) erect head scales not
brilliant white mesally and very dark laterally, 3) mesonotum shortened and arched,
4) mesonotum without broad hoary median longitudinal stripe, 5) acrostichal scal-
ing absent or reduced, 6) coxae more or less same color as adjacent portions of
pleuron, 7) femora and tibiae entirely dark scaled and 8) wing fringe dark; in the
male genitalia by the scaleless sidepieces; as larvae by hair 3-C mesad of 4-C. These
species are widespread in various types of forest in North America. The group con-
sisting of fausti and arboricolus differs as follows: in the adult stage by 1) light
patches on palpus, 2) hindfemur but not hindtibia conspicuously marked with white
and 3) wing with a single light fringe spot at ends of branches of vein R; in the
male genitalia by 1) sparsely scaled sidepiece, 2) clumped spicules on sidepiece and
Zavortink: Treehole Anopheles of New World 3
3) outermost 2 setae on ventral lobe of claspette subequally developed; in the larva
by plumose hairs 11-C, 9-III-VI, 4-IV,V and 10-VII. Both species have restricted
distributions, fausti in the tropical rainforest at the base of the Sierra Madre Orient-
al in central Mexico and arboricolus in the humid montane forest of western Pana-
ma. An. xelajuensis is distinguished as follows: first, by its large size in all stages,
then, in the adult by 1) hindfemur and hindtibia both conspicuously marked with
white, 2) light patch over apex of costa and vein R,, 3) wing with 4 light fringe
spots and 4) wing veins with dark spots caused by accumulation of scales; in the
male genitalia by 1) long sidepiece and 2) numerous scales on sidepiece; in the larva
by 1) hair 3-C thick and blunt, 2) absence of palmate hairs and 3) presence of many
stellate hairs. It is apparently limited to areas of wet montane forest in southern ©
Mexico and Guatemala. An. powderi, unknown in the male and larva, is large in
size, has a single light fringe spot extending from R, to M,4, and has only the
hindfemur conspicuously marked with white. It has been collected in the montane
cloud forest of southern Costa Rica. The differences between these groups are sum-
marized in Table 1. I am not recognizing these groups even informally because the
knowledge of the Central American species is still very fragmentary and I have not
studied the Old World species in detail. |
Very little is known about the bionomics of species in this group. The immature
stages are restricted to treeholes or those artificial containers which approach them
in water composition. Larval development is unusually long. Larvae of barberi are
predacious and those of other species readily eat insect fragments. Adults of both
sexes are occasionally found resting in buildings, culverts and hollow trees and fe-
males of barberi and xelajuensis bite man. Stratman-Thomas and Baker (1936:182-
183) infected barberi with Plasmodium vivax and demonstrated transmission of the
malaria to another person. However, this species has not been incriminated as a nat-
ural vector. American treehole Anopheles extend from the northern United States
(South Dakota to New York) to northern Panama (Chiriqui). All six known spec-
ies are apparently allopatric.
Since An. eiseni Coquillett, 1902, a Neotropical species unrelated to those treat-
ed here, is occasionally found breeding in treeholes, it has been included in the keys.
TAXONOMIC TREATMENT
DESCRIPTION OF GROUP
FEMALES. Head: Adorned with erect scales only; labium entirely dark scaled;
palpus long, about 0.85-1.10 length of proboscis; torus and first flagellar segment
with a few small dark scales. Thorax: Mesonotal bristles quite long; mesonotal scales
absent or restricted to a median tuft on anterior promontory or an acrostichal line:
apn, sp, pra, ppl, stp and upper mep bristles or hairs present, those on stp in a more
or less continuous vertical row; pleuron without scales. Legs: Coxae without scales;
femora and tibiae never mottled, light scales completely absent or restricted to small
patches or rings at apex of segments; tarsi entirely dark scaled; tarsal segment 5 of
all legs much shorter than segment 4. Wing: Never with conspicuous pattern of
light patches or speckles, light scales completely absent or restricted to fringe and
apex of anterior veins. Haltere: Knob dark scaled. Abdomen: Scales completely
absent. Buccopharyngeal Armature: Not studied.
MALES. Similar to females except for sexual differences. Head: Flagellum strong-
ly plumose.
4 Contrib. Amer. Ent. Inst., vol. 5, no. 2, 1969
MALE GENITALIA. Sidepiece: A single strongly developed internal spine and 2
strongly developed parabasal spines present; lateral parabasal spine longer than mes-
al; scales absent, few or numerous. Claspette: Dorsal lobe sclerotized, bearing 3
(rarely 4) overlapping spatulate apically curved setae; ventral lobe conical, spicu-
lose mesally, bearing 3 long and 1 short setae. Clasper: Usually longer than sidepiece.
Phallosome: Aedeagus without leaflets, apex rounded. |
PUPAE. Setae generally weakly to moderately developed, relatively short, fine,
few branched and lightly pigmented. Cephalothorax: Lightly to moderately pig-
mented, with middorsal area, region caudad of trumpet and upper portion of wing-
case slightly darker; hairs 6,7-C relatively close together, separated by a distance
less than 0.5 that between 4,5-C; hair 7-C distinctly smaller than 6-C. Trumpet:
Largely light to dark amber or brown in color, base lighter; rather “‘culicine”’ in
shape, not strongly flared or deeply divided, the pinna relatively small. Abdomen: —
Generally lightly to moderately pigmented, with anterior and/or lateral portions of
most segments slightly darker; hair 9-I small, subequal to or shorter than 3-I, usually
single (single, double); 2-II relatively short, single; 1-III-VII and 5-II,IV frequently
single, never with more than 5 fine branches, less than 0.4 length of corresponding
segment; 5-V-VII similar to 5-III,IV or thickened and then sometimes elongate; 9-
IV-VIII spinelike, long, slender, pointed, becoming progressively longer on poster-
ior segments, usually simple but sometimes with a few coarse branches near tip on
segments VII and VIII. Terminal Segments: Hair 1-[X not irregular as in most other
Anophelini, but instead a simple single or double seta. Paddle: Lightly to moder-
ately pigmented; more or less elliptical to oboval in shape, never more than slight-
ly emarginate; outer part wider than inner; external buttress long; outer margin, to
beyond end of external buttress, serrate, with teeth becoming more prominent and
numerous apically; remainder of margin smooth or weakly serrate, never with long
spicules; hair 1-P thickened, spinelike.
FOURTH INSTAR LARVAE. Head: Setae generally poorly developed; head long-
er and collar narrower than in most Anophelini; maxillary sutures very short; mod-
erately to deeply pigmented, tan to brown, with subantennal area and collar dark-
er and ocular area lighter, never with pattern of contrasting colors; hairs 3-10-C rel-
atively short, fine, usually single (1-4b or f), never large and plumose; 11-C weak-
ly developed, short, usually not extending beyond level of hair 1-A. Antenna: Short,
frequently smooth; uniformly moderately to deeply pigmented, tan to brown, or
becoming lighter apically; hair 1-A short, fine, single, arising on dorsolateral surface
near middle of shaft. Thorax: Integument without conspicuous branched spicules;
hair O-P apparently not developed; 11-P,M,T subequal, very small; 9-M,T not short
and stout, at least as long as hair 10 of corresponding segment; 3-T never palmate.
Abdomen: Hair 1-I not palmate, very small, shorter than hair 2 (except in xela-
juensis); 1-II-VII or 1-II-VU (xelajuensis excepted) palmate, the leaflets well de-
veloped, broad, usually serrate apically; 6-I-VI long, plumose, nearly equally devel-
oped on all segments or 6-III-VI or 6-IV-VI less strongly developed; 4-IV,V (except in
xelajuensis) elongate, longest hair on dorsal surface of respective segment; 5-IV-VII
(except in xelajuensis) elongate, with relatively few weak and short lateral branches.
Spiracular Lobe: Pecten with most teeth, except apically, long, never with regular
sequence of long and short teeth; hair 1-S usually single (1-3b) or with fine apical
branches, never with numerous long branches from base.
zit).
3(2).
4(2).
5(4).
Zavortink: Treehole Anopheles of New World 5
KEYS TO SPECIES
ADULTS
Hindtibia, but not hindfemur, with a conspicuous broad apical white ring;
vein R sometimes with light patches at level of ends of veins 1A and Rs
‘(see Systematics). . . icee 2 CEST
Hindtibia without a conspicuous broad apical white ring, apex dark or with
a conspicuous white patch; hindfemur sometimes with a conspicuous api-
cal white ring; vein R without light patches at level of ends of veins 1A
OU Fe a ag wid ew te ae eae ek Ga a Mi ae
Mesonotum without a broad hoary median longitudinal stripe and with scales
absent or restricted to a tuft in center of anterior promontory; hindfemur
entirely dark scaled: wing fringe entirely dark scaled . ..°..... . «0
Mesonotum with a broad hoary median longitudinal stripe and a long acros-
tichal scale line; hindfemur with apical white ring; wing fringe with api-
cal light spot
Bristles on ppl 6-11; anterior forecoxal bristles 19-32; anterior acrostichal
bristles dark and usually not accompanied by scales; light erect head scales
dingy yellowish (fig.2). . . . . . .L. barberi
Bristles on ppl 2-5; anterior forecoxal bristles be 18: anterior acrostichal brist-
les amber and apoommpanied by 5-20 whitish scales: light erect head scales
POM Wey barteri! dhe ula 6 we Cs 1 i ee
Apex of costa and vein R, light scaled; small light fringe spots present at
ends of veins i i, 44 and Cu, ; dark wing spots present (fig. 12).
ogee xelajuensis
Apex of costa and vein | Bi dak scaled: without ‘frinee spots at ends of veins
Mi42,M3+44 and Cu, ; wing without dark BOOTS. ah i Ey -4 Gee. ce
Palpus with light scales at joints and apex; wing moderately scaled; scales in
middle of vein 1A delish vein or slightly spreading (fig. 12). . .
. . 3. fausti; 4. arboricolus
Palpus entirely ‘dark scaled: wing profusely scaled: scales in middle of vein
1A wide spreading (fig. ee jk pu inch ge deities 0 bial Achebe iat i
MALE GENITALIA
(6. powderi unknown)
_ Aedeagus with 1 pair of conspicuous serrate leaflets; dorsal claspette lobe
with 2 terete setae (see Systematics) . . cite ck AIST
Aedeagus without leaflets; dorsal claspette lobe with z flattened SoS © | ave
Sidepiece long, scales numerous (fig.9) . .... . . . . 5.xelajuensis
Sideniece shorter, 6cales abeont Op Gawe io os ely pith aint cath eA | ult
Lateral 2 setae of ventral lobe of claspette subequal in development; spicules
of sidepiece grouped into large clumps; sidepiece usually with at least 1 or
4(3).
2(1).
seh
4(3).
3(2).
Contrib. Amer. Ent. Inst., vol. 5, no. 2, 1969
2 Beales (igs 2) os ak . . . . 3. fausti; 4. arboricolus
Lateral seta of ventral lobe of claspette not as strongly developed as adjacent
seta; spicules of sidepiece arranged singly, in rows, or in small clumps; side-
PieOe VSURIy WIKROUIE SCAMS Gs hee re ee ee St
Parabasal spines much more strongly developed than internal spine; lateral .
parabasal spine flattened preapically; sidepiece usually more or less ellip-
tical in shape Cie Ss . . . .L. barberi
Parabasal spines only slightly stronger than internal spine: lateral parabasal
spine not flattened preapically; sidepiece usually more or less cylindrical
or sort comical in Shape (fig. Ss a ee er ee 2 ithe
PUPAE
(3. fausti and 4. arboricolus unknown)
Trumpet deeply divided; paddle margin with fringe of ie fine spicules (see
Systematics). . . re ay
Trumpet not deeply divided: - paddle margin without fringe . Sade its Gu
Hair 7-VII single, longer than 9-VIII; hair O-VII enlarged, multiple; 5-V more
strongly developed than 1-V (fig. 9). Bint . . 9. xelajuensis
Hair 7-VII 2,3b (1-4), shorter than 9-VIII; hait 0-VII small, — —
double); 5 VV usually less strongly developed than 1-V.. . a
Large species, width of segment VIII greater than 1.00 mm (fig. 11) .
ee Ey ats A er ph a DR OS Ow er
Small species, width of segment VIII lessthanO.85mm....... .4
Hair 9-III-VIII darkly pigmented; 9-III more similar in size to 9-IV than 9-II;
hair 5-VI,VII thickened, single (fig.3). . . pak, barberi
Hair 9-ITI-VIII concolorous with integument; 9. Il intermediate in size be-
tween 9-II and 9-IV; hair 5-VI and rsp 5-VII fine, 2,3b (fig. 5) .
. 2. Judithae
FOURTH INSTAR LARVAE
(6. powderi unknown)
Hairs 4-6-C long, plumose; 3-T palmate (see Systematics) . . . . . eiseni
Hairs 4-6-C shorter, never plumose; 3-T not palmate .. .°. . 2.1. . 42
Hair 1-III-VII stellate; 9-III-VI very strongly developed, stellate, long, 6,7b (5-
9); hair 3-C thickened, apex usually blunt (fig. 10). . . . 5. xelajuensis
Hair 1-III-VI palmate; 9-III-VI moderately developed, not stellate, elongate
and plumose or shorter and 2-4b (1-5); hair 3-C fine, apex attenuate . .3
Hair 3-C mesad of 4-C; hair 9-III-VI not elongate or plumose; 7-VIJ much
more strongly developed than GV | .4
Hair 3-C laterad of 4-C; hair 9-ITI-VI iepantly plumose TNIL not a as strongly
developed as'6-VIF coo 4 igi alae Cem
Zavortink: Treehole Anopheles of New World 7
4(3). Inner clypeals (2-C) widely spaced, separated by a distance greater than 2.0
that between inner and outer clypeals (3-C); hair 13-II-V,VII usually 3b (3-
5); integument on underside of prothorax and abdominal segments I-VIII
spiculose (fig.4) . . . . . 1. barberi
Inner clypeals (2-C) closely approximated, evan by a distance less than
that between inner and outer clypeals (3-C); hair 13-II-V,VII usually single
(1-3b); integument of thorax and abdomen not spiculose (fig. 6)
eas judithae
5(3). Hair 9-LII elongate, plumose; 13-II-V with many fine inconspicuous branches
beyond middle; antennal shaft with conspicuous spicules (fig. 7) ‘et
. . &. fausti
Hait 9 J I not elongate or - plumose; 13 TLV with 3 4 (2- 6) conspicuous
branches from base; antennal shaft without conspicuous spicules (fig. 8)
. 4. arboricolus
1. Anopheles (Anopheles) barberi Coquillett
Figs. 1-4
1903. Anopheles barberi Coquillett, 1903:310. TYPE: Holotype ?, Plummer’s Island, Maryland,
United States, 17 Aug 1903, H.S. Barber [USNM, 6959].
Anopheles (Anopheles) barberi of Edwards (1921:272); Dyar (1928:454); Matheson (1944:114-
115); Darsie (1949:524-525); Penn (1949:65-66); Carpenter and La Casse (1955:32-34, in
part); Stone, Knight and Starcke (1959:15, in part); Carpenter (1968:72, in part); Zavortink
(1969:31-33).
Anopheles barberi of Dyar (1904:243-244); Dyar and Knab (1907:49); Stratman-Thomas and
Baker (1936:182-183); Vargas (1942a:172,173,174; 1942b:329-331); Russel, Rozeboom and
Stone (1943:21,31, in part); Jenkins and Carpenter (1946:35-36, in part); Petersen, Chapman
and Willis (1969:134-135).
Anopheles (Coelodiazesis) barberi of Dyar (1918:142); Vargas (1943:64,65 ,66,67, in part); Vargas
and Martinez (1956:111-113,140, in part); Vargas (1959:373).
Coelodiazesis barberi of Dyar and Knab (1906:177); Howard, Dyar and Knab (1917:1036-1038).
FEMALE (fig. 2). Wing: 3.49 mm. Proboscis: 1.73 mm. Forefemur: 2.10 mm.
Abdomen: about 2.2 mm. A small light-colored species. Head: Integument light to
dark brown; interocular bristles dark or amber; erect scales long, usually dingy yel-
lowish mesally, dark or dingy yellowish laterally; interocular scales absent or very
few, short and dingy yellowish; palpus entirely dark scaled, the proximal scales ap-
pressed. Thorax: Mesonotum shortened and arched; mesonotal integument very
light to dark brown, largely shining, without a broad hoary median longitudinal
stripe; pleural integument lighter than that of mesonotum or becoming lighter ven-
trally; mesonotal bristles numerous, strongly developed, very long and conspicuous,
dark in color; mesonotum without scales or sometimes center of anterior promon-
tory with 1-4 narrow dark scales; ppl bristles 6-11. Legs: Coxae more or less same
color as adjacent portions of pleuron; anterior forecoxal bristles 19-32; femora, tib-
iae and tarsi entirely dark scaled. Wing: Veins and fringe entirely dark scaled, scales
on veins uniformly distributed and not grouped into dark spots; moderately scaled;
scales in central portion of vein 1A clasping vein or slightly spreading. Abdomen:
8 Contrib. Amer. Ent. Inst., vol. 5, no. 2, 1969
Integument of tergites light to dark brown, usually somewhat dappled, sternites
lighter.
MALE (fig. 2). As for female except for sexual differences.
MALE GENITALIA (fig. 3). Sidepiece: More or less elliptical in shape, short;
spicules arranged singly, in rows, or in small clumps; scales usually absent; para-
basal spines much more strongly developed than internal spine; lateral parabasal —
spine flattened preapically, apex sharply attenuate and recurved or sinuous; inter-
nal spine not more strongly developed than large setae of sidepiece, apex usually
sinuous. Claspette: Most lateral seta of ventral lobe distinctly shorter and/or finer
than seta next mesad. Clasper: Without spicules near base.
PUPA (fig. 3). Abdomen: 3.04 mm. Trumpet: 0.37 mm. Paddle: 0.88 mm. Width
of segment VIII: 0.73 mm. Cephalothorax: Moderately pigmented with darker areas
more extensive than in judithae; hairs 1,3-C subequal in length. Trumpet: Dark am-
ber to brown in color. Abdomen: Moderately pigmented, darker areas more exten-
sive than in judithae; hairs 6,7-I subequal in length; 1-II,III usually single or double;
5-III-V short, fine, usually single; 5-VI,VII thickened and single, usually shorter than
to subequal in length to hair 9 of corresponding segment; 9-III-VIII darkly pigment-
ed, strongly contrasting with abdominal integument; 9-III approaching 9-IV in both
diameter and length; O-VII very small, usually single (single, double); 7-VII usually
double (single, double), shorter than 9-VIII. Paddle: Moderately pigmented, midrib
darker.
FOURTH INSTAR LARVA (fig. 4). Head: 0.65 mm. Anal Saddle: 0.27 mm.
Head: Inner clypeals (2-C) widely spaced, usually separated by a distance greater
than 2.0 that between inner and outer clypeals, single and simple; outer clypeals
(3-C) far mesad of 4-C, fine, single; 11-C not plumose, single or 2,3b. Antenna:
Spicules absent or inconspicuous; hair 4-A simple and single. Thorax: Integument,
especially on underside of prothorax, spiculose; hair 2-P short, plumose, with num-
erous lateral branches; 8-P long, shaft not notably thickened, lateral branches short
and moderately numerous; 8-M short, all lateral branches long; 8-T a normal plu-
mose hair; 9-P,M,T spiculate. Abdomen: Integument on underside of segments I-
VIII spiculose; hair 1-I very short, single, 1-II simple to palmate, 1-III-VII palmate;
6-I-VI similar in development and length; 9-I-VI moderately developed, with branch-
es arising near base, never stellate or plumose; 13-II-V,VII moderately developed,
usually 3b (3-5); hair 4-IV,V simple, single; 5-VII moderately long with 2,3b (1-3)
arising near base; 7-VII moderately long, usually single or double (1-3b); hair 10-
VII not plumose but single or 2,3b or 2,3f. Segment VIII: Hairs 3,5 usually with
3,4b (1-5) arising near base. Anal Segment: Spicules of saddle moderately conspic-
uous, located along ventral and caudal margins; hair 1 simple, usually single.
SYSTEMATICS. Although barberi and judithae are a pair of closely related allo-
patric species, they are easily separated in all stages on the basis of the characters
given in the keys.
Examination of more specimens since the publication of my earlier paper (Zavor-
tink, 1969) has indicated that some changes need to be made in the description of
barberi presented there. In the male genitalia the ratio of the distance from the para-
basal spines to the internal spine to the distance from the latter to the apex of the
sidepiece should be broadened from 0.8-1.0 to 0.6-1.6 and the entire proctiger sec-
tion should be deleted. In the pupa hair 5-VI,VII is sometimes longer than previ-
ously indicated and is then subequal in length to hair 9 of the corresponding seg-
ment. In the larva hair 1-II is frequently palmate and 7-VII is often double or triple.
The following extremes in variation of chaetotaxy of the pupa have been seen:
Zavortink: Treehole Anopheles of New World 9
hair 5-VI is fine, as in judithae, on a single specimen from Ithaca, New York; hair
5-V is thickened, as in xelajuensis, on one side of the only specimen available for
study from the Baltimore, Maryland, area; hair 9-VI,VII is shortened in some spec-
imens from Ithaca and hair 5 of the corresponding segment is then longer than it.
An. barberi is apparently the most modern species of the group. It is widespread,
occurring throughout the eastern half of the United States, and the vestiture of the
adult is the most derived of any New World treehole Anopheles.
BIONOMICS. An. barberi is found at elevations of less than 1200 meters. The
immature stages are found in rot holes in trees and stumps and in artificial con-
tainers, particularly those made of wood or containing leaves and twigs. Since long
periods of unfavorable weather, at least in the northern portion of its range, are
passed in the larval stage, the species is usually recovered from sites which contain
water more or less continuously. More than a dozen larvae are seldom collected to-
gether; this is apparently due to their predacious nature. Adults are occasionally
found resting in buildings and culverts and under bridges. Both sexes are attracted
to lights and females are attracted to humans.
DISTRIBUTION (fig. 1). An. barberi is the most widely distributed of the New
World treehole Anopheles, being found at low to moderate elevations throughout
the eastern United States, from South Dakota and New York in the north to Texas
and Florida in the south. Material examined: 239 specimens; 60 males, 19 male gen-
italia, 68 females, 1 adult, 34 pupae, 57 larvae; 24 individual rearings (3 pupal, 16
larval, 5 incomplete).
UNITED STATES. Alabama: Guntersville Lake, 28 May-27 Aug 1942, 3 9? [UCLA]. Ozark,
Camp Rucker, 11 Mar 1943, J.G. Franclemont, 1 ¢ [CU] . Sheffield, 18 Aug 1942, J.N. Belkin, 1 ?
[UCLA]. Wilson Dam, 10 June-19 July 1942, J.N. Belkin, 1 lp? (167), 1 pd (171), 1 p? (168), 2
6, 2 6 gen [UCLA]; 1942, 1 ¢6 [HOPK]. Arkansas: Little Rock, Little Fourche Bayou, 28-29 Mar
1943, J.N. Belkin, 2 lp (440,448), 3 P [UCLA]. Scott, 2 Oct 1908, J.K. Thibault, 1 d, 1 d gen, 1
9 [USNM] . Delaware: Bombay Hook, 17 June 1964, R.W. Lake, 1 L [USNM]; 19 Aug 1965, R.W.
Lake, 1 9 [USNM]. Cooch’s Bay, 26 June 1961, R.W. Lake (224), 1 6, 1 d gen [USNM] . District
_ of Columbia: Kenilworth, 2 July 1943, C.W. Travis, 1 9 [USNM]. Soldier’s Home, 3 July 1943,
N.E. Good and C.W. Travis, 2 6, 1 6 gen [USNM]. No locality, 12 Aug 1944, N.E. Good, 1 L
[USNM]. Georgia: Atlanta, Fort McPherson, 19 Apr 1943, 2 L [USNM]. Milledgeville, 14 June
1950, R.H. Foote, 1 L [USNM]. Savannah, Chatham Field, 17 Aug 1944, 2 L [USNM] . Lowa:
Ames, 10 Sept 1919, Bishopp (9085), 2 6, 1 6 gen [USNM]. Oskaloosa, 14 Sept 1933, 1 2 [US-
NM]. Kentucky: Louisville, GE. Quinby, 1 6 [USNM]. Maxon Mill, 17 June 1935, G.E. Quinby,
1 6, 1 6 gen [USNM]. Louisiana: Alexandria, Esler Field, 13 Oct 1942, W.W. Wirth, 1 ¢d [USNM].
Baton Rouge, 1941-1947, W.W. Wirth, 2 6, 1 ¢ gen, 1 ?, 1 L [USNM]. Lake Charles, 15 June
1943, W.W. Wirth, 1 ¢ [USNM]. Olla, May 1943, 1 9 [USNM] . Maryland: Annapolis, 6 July 1933,
F.C. Bishopp, 1 9 [USNM]. Baltimore, July 1931, 1 L [HOPK]. Baltimore, Gwynns Falls Park,
July 1931, 1 lp [HOPK]. Baltimore, Patapsco State Park, 17 Oct 1965, W.A. McDonald (UCLA
286), 1 L [UCLA]. Bethesda, 22 Aug 1944, G.B. Vogt, 1 6, 1 9 [USNM]. Cabin John, Aug-Oct
1908, F. Knab, 3 6, 5 9, 2 L [USNM]; July 1965, W.A. McDonald (UCLA 286A), 1 L [UCLA].
Great Falls, 18 May 1919, W.L. McAtee, 1 9 [USNM]. Plummer’s Island, 23 Aug-10 Sept 1903,
H.S. Barber, 2 9 [USNM]; 5 Sept 1904, H.S. Barber and E.A. Schwarz, 1 ? [USNM]; 10-13 July
1905, HS. Barber, 2 6, 2 6 gen, 4 9 [USNM]; 30 Aug 1908, F. Knab, 1 d6 [USNM]; 24 May
1912, E.A. Schwarz and H.S. Barber, 1 9? [USNM]; 19 June 1912, H.S. Barber, 1 6 [USNM]; 13
Sept (10263), eggs, 3 L [USNM]; 1 9 [USNM]. Mississippi: Agricultural College, 23 July-15 Oct
1905, G.W. Herrick, 2 2 [USNM]. Clinton, 30 May 1945, 1 d [USNM]. Greenwood, 1928, 2 d, 1
é gen, 3 9 [USNM]. Hattiesburg, Camp Shelby, 22 June 1943, 2 L [USNM]. Missouri: Neosho,
Camp Crowder, 20 July 1942, A.B. Gurney (96), 1 2, 1 P [USNM]; Sept 1942, A.B. Gurney
(196), 19, 1P,1L [USNM]. St. Louis, Aug, A. Busck, 1 9 [USNM] . New Jersey: Chester, 8 Sept,
J.M. Aldrich, 1 9 [USNM]. New York: Ithaca, 15 June-15 Oct 1932, R. Matheson, 5 d, 10, 1 Ip
(1027-1065), 5 P, 8 L [CU], 2 6, 1 6 gen, 3 9 [UCLA]; 1933, 1 L [CU]; 20 Aug 1935,
10 Contrib. Amer. Ent. Inst., vol. 5, no. 2, 1969
16,1 A [CU]. North Carolina: Fayetteville, Fort Bragg, 1 Aug 1945, 1 ? [USNM]. Montreat, 25
June 1940, 1 9 [HOPK]. Ohio: Canton, 22 Dec 1967, T.J. Zavortink (UCLA 437), 10 lpd (437-
30-36,39,41,43), 4 lp? (437-37,38,40,42), 1 pd (437-44), 2 d gen, 1 L [UCLA] . Oklahoma: Ida-
bel, 7 June 1938, L.E. Rozeboom, 1 lp (A-42) [HOPK]. South Carolina: Charleston, Charleston
Field, 17 Aug 1944, 2 L [USNM]. Columbia, 1 Aug 1906, 1 6, 1 dé gen, 1 9 [USNM]. Tennessee:
Kentucky Lake, 26 Aug 1942, 1 6, 1 6 gen [UCLA]. Paris, Camp Tyson, 21 June 1944, 2d
[USNM]. Reelfoot Lake, 11 Sept 1939, T.W. Simpson and G.E. Quinby, 1 L [HOPK]. Virginia:
Bluemont, 29 July 1904, 1 d [USNM]. Dead Run, 2 July, R.C. Shannon, 3 6, 1 d gen [USNM].
Newington, 22 Aug 1910, S.A. Rohwer, 1 2 [USNM]. Williamsburg, Camp Peary, July 1943,R.
Bohart, 1 L [USNM]. Woodstock, 9 Aug 1904, F.C. Pratt, 2 6, 1 9? [USNM]. Texas: Abilene, 30
Aug 1953, R.X. Schick, 2 9 [UCLA]. No Data: (Mound 10180), 1 Ip? (10180A3), 1 d, 2 ¢
[USNM]; (Mound 10253-5), 1 6 gen [USNM]; E.B. Johnson (37-268), 1 ¢ [USNM] ; (57-3), 1 L
[USNM]; (125-1), 1 2 [USNM]; (140-2), 1 @ [USNM]; (150-3), 1 ¢ gen, 1 L [USNM] ; (150-4),
1 6 [USNM]; 1 2? [USNM];1L [CU].
2. Anopheles (Anopheles) judithae Zavortink
Figs. 1,5,6
1969. Anopheles (Anopheles) judithae Zavortink, 1969:28-31. TYPE: Holotype 6 with associ-
ated larval and pupal skins (UCLA 302-40), Cochise Stronghold Recreation Area, Dragoon
Mountains, Cochise County, Arizona, United States, larva from oak treehole, 22 Mar
1966, T.J. Zavortink [USNM].
Anopheles (Anopheles) barberi of Vargas (1940:319-322); Carpenter and La Casse (1955:32-34,
in part); Stone, Knight and Starcke (1959:15, in part); Carpenter (1968:72, in part).
Anopheles barberi of Russel, Rozeboom and Stone (1943:21,31, in part); Jenkins and Carpenter
(1946:35-36, in part); Richards, Nielsen and Rees (1956:14); Rigby, Blakeslee and Forehand
(1963:50); Burger (1965:396); Nielsen, Arnell and Linam (1967:76); Nielsen, Linam, Arnell
and Zavortink (1968:363). :
Anopheles (Coelodiazesis) barberi of Vargas (1943:64,65,66,67, in part); Vargas and Martinez
(1956:111-113,140, in part).
FEMALE. Wing: 3.31 mm. Proboscis: 1.92 mm. Forefemur: 2.18 mm. Abdomen:
about 2.1 mm. Very similar to barberi but differing in the following. Head: Inter-
ocular bristles amber; erect scales long, entirely pale white or some lateral ones dark;
interocular scales numerous, white, usually none appreciably elongate. Thorax: An-
terior acrostichal bristles amber; mesonotal scaling restricted to a small tuft of 5-20
whitish scales in center of anterior promontory; ppl bristles 2-5. Legs: Anterior fore-
coxal bristles 6-18.
MALE. Similar to female except for sexual differences.
MALE GENITALIA (fig. 5). Sidepiece: More or less cylindrical or short conical
in shape, short; spicules arranged singly, in rows, or in small clumps; scales absent;
parabasal spines only slightly more strongly developed than internal spine; lateral
parabasal spine not flattened preapically, apex gradually attenuate and strongly re-
curved; internal spine more strongly developed than large setae of sidepiece, apex
usually recurved. Claspette: Most lateral seta of ventral lobe distinctly shorter and/or
finer than seta next mesad. Clasper: Without spicules near base.
PUPA (fig. 5). Abdomen: 2.80 mm. Trumpet: 0.37 mm. Paddle: 0.83 mm. Width
of segment VIII: 0.75 mm. Cephalothorax: Usually lightly pigmented with darker
areas less extensive than in barberi; hairs 1,3-C subequal in length. Trumpet: Light
Zavortink: Treehole Anopheles of New World it
amber in color. Abdomen: Usually lightly pigmented, darker areas less extensive
than in barberi; hairs 6,7-I more or less subequal in length; 1-II,III usually single
(1-3b); hair 5-III-VI short, fine, usually 2,3b; hair 5-VII usually short, fine and 2,3b,
sometimes thickened and single, but then longer than 9-VII; hair 9-III-VIII lightly
pigmented, concolorous with abdominal integument; 9-III more or less intermediate
in diameter and length between 9-II and 9-IV; hair O-VII small, single; 7-VII usually
2,3b (1-4), shorter than 9-VIII. Paddle: Usually lightly pigmented, midrib darker.
FOURTH INSTAR LARVA (fig. 6). Head: 0.67 mm. Anal Saddle: 0.30 mm.
Head: Inner clypeals (2-C) closely approximated, separated by a distance less than
that between inner and outer clypeals, single and simple; outer clypeals (3-C) far
mesad of 4-C, fine, double; 11-C not plumose, single or 2,3b. Antenna: Spicules
inconspicuous; hair 4-A simple, forked apically. Thorax: Integument without spic-
ules; hair 2-P moderately long, with only a few (2-6) long lateral branches; 8-P mod-
erately long, shaft not conspicuously thickened, lateral branches long and moder-
ately numerous; 8-M moderately long, apical lateral branches very long; 8-T a nor-
mal plumose hair; 9-P,M,T simple. Abdomen: Integument without spicules; 1-I very
short, usually single (1-3b), hair 1-II-VII palmate; 6-III-VI less strongly plumose than
6-I,I]; hair 9-I-VI moderately developed, most branches arising near base, never stel-
late or plumose; 13-II-V,VII moderately developed, usually single (1-3b); hair 4-IV,V
simple, single; 5-VII long with 3,4b arising from near base; 7-VII moderately long
with 2,3b arising from near base; 10-VII not plumose, but with 2,3b arising from
near base. Segment VIII: Hair 3 with 2-4b arising from base; 5 usually single. Anal
Segment: Spicules of saddle inconspicuous, located along caudal margin and distal
half of ventral margin; hair 1 simple, single.
SYSTEMATICS. An. judithae is the treehole Anopheles of the southwestern Unit-
ed States. Although it is quite distinct from barberi in all stages, it was confused
with that species for nearly 30 years. The two can be easily distinguished on the
basis of the diagnostic features given in the Keys.
Although pupal hair 5-VII is fine and 2,3b in most specimens of judithae, it is
thickened, single and elongate in a few individuals from nearly every collection.
This is true not only in the population in the Gila River system, but also in the one
in the Chisos Mountains of Texas.
BIONOMICS. An. judithae occurs at elevations between 600 and 2200 meters.
At the upper limit of its altitudinal range it occurs in the more or less continuous
xeric evergreen forest, but at lower elevations it is restricted to the narrow bands
of trees lining watercourses. The immature stages have been taken only from tree-
holes. Like barberi, judithae overwinters in the larval stage and, as a result, is more
frequently found in permanent than temporary treeholes. In contrast to barberi, it
is not unusual to find great numbers of judithae larvae in the same treehole. Except
for 3 males collected in a building in Sonora, Mexico (Vargas 1940:319-322), adults
have not been encountered outside the laboratory.
DISTRIBUTION (fig. 1). An. judithae is widely distributed at moderate to high
elevations in the southwestern United States and, undoubtedly, northern Mexico.
In Arizona and western New Mexico it is common in the Gila River system. In
Texas it has been collected in the southern Trans-Pecos area. Material examined:
1002 specimens; 200 males, 15 male genitalia, 190 females, 304 pupae, 293 larvae;
168 individual rearings (56 pupal, 102 larval, 10 incomplete).
MEXICO. Sonora: Imuris, 1940, A. Martinez Palacios, 1 6,16 gen [ISET].
UNITED STATES. Arizona: Chiricahua National Monument, Headquarters, 24 Dec 1966, L.T.
Nielsen (N-36-66), 1 6 [UTAH] ; same data (N-37-66), 3 6, 4 9 [UTAH] . Cochise Stronghold Rec-
f2 Contrib. Amer. Ent. Inst., vol. 5, no. 2, 1969
reation Area (Dragoon Mts.), 22 Mar 1966, T.J. Zavortink (UCLA 302), holotype lpdé (302-40),
allotype lp? (302-42) [USNM], 6 lpd (302-25 ,41,43,45 46,48), 5 Ip? (302-44,47,49,51,52), 3 pd
(302-100,101,103), 1 p? (302-102), 1 9, 1 P, 6 L [UCLA] ; 23 Mar 1966, T.J. Zavortink (UCLA
306), 1 Ip? (306-20) [UCLA] ; 4 Sept 1966, T.J. Zavortink (UCLA 328), 1 lpd (328-22) [UCLA] ;
6 Sept 1966, T.J. Zavortink (UCLA 342), 7 Ipd (342-13,15 30,3338 43,44), 11 lp? (342-10,31,
3234-37 39-42), 10 pd (342-100-106,109,110,113), 5 p? (342-107,108,111,112,114),17d,2¢
gen, 25 9°, 48 P, 27 L [UCLA]. Coronado National Memorial, Headquarters, 20 Mar 1968, L.T.
Nielsen, J H. Arnell and J.H. Linam (HA-10-68), 2 6, 2 9 [UTAH] . Coronado National Memorial
(1 mi E), 20 Mar 1968, L.T. Nielsen, J H. Arnell and J.H. Linam (HA-9-68), 2 6, 1 9 [UTAH].
Douglas (34 mi ENE), 13 Sept 1968, L.T. Nielsen (N-43-68), 2 2, 2 L [UTAH]. Fort Huachuca,
17 Sept 1962, Bates, 1 d gen [USNM]. Kitt Peak National Observatory (Quinlan Mts.), 12 Sept
1968, T.J. Zavortink (UCLA 448), 1 Ipd (448-27), 6 Ip? (448-20-25), 1 p? (448-15), 1 lp (448-28)
[UCLA] ; 8 Sept 1969, T.J. Zavortink (UCLA 630), 1 pd (630-103), 1 P, 14 L [UCLA]. Lochiel
(2 mi E), 21 Mar 1968, L.T. Nielsen, J H. Arnell and J.H. Linam (HA-13-68), 1 6 [UTAH].
Lochiel (11 mi E), 21 Mar 1968, L.T. Nielsen, J H. Arnell and J.H. Linam (HA-12A-68), 4 L
[UTAH]. Nogales (13 mi NW), 21 Mar 1966, T.J. Zavortink (UCLA 297), 2 Ip? (297-20,21)
[UCLA] . Nogales (9 mi NE), 21 Mar 1966, T.J. Zavortink (UCLA 298), 3 Ipd (298-40,42,49), 4
Ip? (298-43,45 50,51), 3 pd (298-101-103), 4 lp (298-41 ,46-48), 1 d gen, 3 9,4 P, 22 L [UCLA];
same data (UCLA 299), 5 Ipd (299-21 ,26,28-30), 6 lp? (299-20,23-25 27,31), 1 lp (299-22),1¢
gen, 2 L [UCLA]. Palominas (1 mi E), 20 Mar 1968, L.T. Nielsen, J H. Arnell and J.H. Linam
(HA-8-68), 1 6, 1 9 [UTAH]. Patagonia (1-4 mi WSW), 24 Aug 1954, W.A. McDonald (UCLA
137), 2 L [UCLA]; 6 Sept 1963, J. Burger (UCLA 413A), 2 L [UCLA]; 18 Aug 1964, J. Burger
(UCLA 253), 17 3, 11 9 [UCLA]; 13 Sept 1964, J. Burger (UCLA 270), 3 L [UCLA]; 20 Sept
1964, J. Burger (UCLA 260), 17 6,1 d gen, 9 9 [UCLA]; 25 July 1965, J. Burger (UCLA 281), 4
L [UCLA]; 27 July 1965, J. Burger (UCLA 282), 5 L [UCLA]; 21 Mar 1966, T.J. Zavortink
(UCLA 300), 2 Ipd (300-30,38), 6 Ip? (300-32-36,39), 2 pd (300-100,102), 2 p? (300-101,103), 3
6,3 2,6 P, 12 L [UCLA]; 5 Sept 1966, T.J. Zavortink (UCLA 333), 1 6,1 P,2 L [UCLA]; same
data (UCLA 334), 1 Ipd (334-15), 4 Ip? (334-12-14,16), 2 pd (334-102,103), 3 p? (334-100,101,
104), 1 9,3 P, 2 L [UCLA]; 21 Mar 1968, L.T. Nielsen, J H. Arnell and J.H. Linam (HA-14-68),
18 d, 17 2, 11 L [UTAH]; 14 Sept 1968, T.J. Zavortink (UCLA 458), 1 p? (458-100). Portal (3
mi W), 20 Mar 1968, L.T. Nielsen, J H. Arnell and J.H. Linam (HA-7A-68), 1 6 [UTAH]. Portal
(15 mi WNW), 6 Sept 1966, T.J. Zavortink (UCLA 343), 9 Ipd (343-14,15,17,31-34,36,37), 8 lp?
(343-12,16,19,30,38-41), 10 pd (343-100-104,107,109,110,113,114), 4 p? (343-105,108,111,
112), 2 Ip (343-13,35), 30 6, 2 3 gen, 25 9, 69 P, 39 L [UCLA]. Prescott, 19 Mar 1966, L.T.
Nielsen (N-2-66), 2 6, 1 9, 1 P, 5 L [UTAH]. Prescott (7 mi NE), 1 Sept 1966, T.J. Zavortink
(UCLA 315), 2 Ipd (315-16,18), 1 6 gen [UCLA]. Santa Rita Mts., 20 Oct 1940, R.A. Flock, 1 ¢,
1 2 [CU], 1 6, 1 d gen [UCLA]. Wickenburg (5 mi S), 18 Mar 1966, L.T. Nielsen (N-1-66), 1 6,
1 d gen, 1 9,9 L [UTAH] . New Mexico: Animas (15 miS), 19 Mar 1968, L.T. Nielsen, J H. Arnell
and J.H. Linam (HA-6C-68), 1 9? [UTAH]. Glenwood (5 mi E), Catwalk Cmpg., 21 Mar 1967, L.T.
Nielsen and J.H. Linam (N-5-67), 1 6, 1 6 gen, 1 P, 1 L [UTAH]. Texas: Big Bend National Park,
Chisos Mts., 31 Aug 1969, T.J. Zavortink and J.A. Bergland (UCLA 603), 1 lpd (603-23), 4 lp?
(603-21 ,22,26,33), 6 pd (603-100,101,102,103,108,109), 2 p? (603-104,110), 2 lp (603-20,27), 2
d gen, 1 P, 4 L [UCLA]; 1 Sept 1969, T.J. Zavortink (UCLA 612), 2 lpé (612-10,11), 2 lp? (612-
20,24), 1 L [UCLA] ; same data (UCLA 613), 1 lpd (613-14) [UCLA].
3. Anopheles (Anopheles) fausti Vargas
Figs..1/ 42
1943. Anopheles (Coelodiazesis) fausti Vargas, 1943:66-68. TYPE: Holotype larva, Tamazun-
chale, San Luis Potosi, Mexico, from treehole, Apr or May 1942, M. Macias [ISET].
Anopheles (Anopheles) fausti of Stone, Knight and Starcke (1959:18, in part); Belkin, Schick and
Zavortink: Treehole Anopheles of New World 13
Heinemann (1965 :34); Zavortink (1969:30, in part).
Anopheles (Coelodiazesis) fausti of Vargas and Martinez (1956:113-116,141); Senevet (1958:43);
Vargas (1959:377).
Anopheles xelajuensis of Vargas (1942a:169-175); Russel, Rozeboom and Stone (1943:35, in
part).
FEMALE (fig. 12). Wing: 3.36 mm. Proboscis: 2.06 mm. Forefemur: 1.98 mm.
Abdomen: about 2.0 mm. A small dark species. Head: Integument dark brown; in-
terocular bristles white; erect scales apparently short, dark brown except for a small
patch of bright white scales in anterior dorsal region; interocular scales very long,
fine, wavy, white, forming a conspicuous tuft; palpus with white scales at joints and
apex, proximal scales slightly outstanding. Thorax: Mesonotum elongate and rather
flat; mesonotal integument dark brown except for broad hoary median longitudinal
stripe; pleural integument largely dark brown; mesonotal bristles not as long, strong
or conspicuous as in barberi and judithae, dark except for light anterior acrostichals;
center of anterior promontory with a conspicuous tuft of white scales; acrostichal
scale line extending about halfway to scutellum; pp/ bristles 3-5. Legs: Coxae and
trochanters white, strongly contrasting with dark pleuron; anterior forecoxal bris-
tles 13-20; femora, tibiae and tarsi dark scaled except as follows: apex of fore- and
midfemora with a few white scales, more conspicuous on midfemur; apex of hind-
femur with a broad ring of semierect white scales; apex of tibiae usually with a few
white scales. Wing: Dark scaled except for a single large cream-colored apical fringe
spot at ends of branches of vein R; scales of veins uniformly distributed and not
grouped into dark spots; moderately scaled; scales in middle of vein 1A nearly clasp-
ing vein. Abdomen: Tergites and sternites dark brown.
MALE. As for female except for usual sexual differences.
MALE GENITALIA (fig. 12). Sidepiece: More or less cylindrical or short conical
in shape, short; spicules grouped into large clumps; 1 to 5 scales usually present;
parabasal spines slightly to considerably stronger than internal spine; lateral para-
basal spine flattened preapically, apex attenuate and straight to recurved; internal
spine as or more strongly developed than large setae of sidepiece, apex nearly
straight, curved or recurved. Claspette: Lateral 2 setae of ventral lobe subequal in
length and stoutness. Clasper: Sometimes with a few spicules near base of ventral
surface.
PUPA. Unknown.
FOURTH INSTAR LARVA (fig. 7). Head: 0.64 mm. Anal Saddle: 0.27 mm.
Head: Inner clypeals (2-C) closely approximated, separated by a distance much less
than that between inner and outer clypeals, single, weakly plumose at least apically;
outer clypeals (3-C) distinctly laterad of 4-C, fine, usually forked apically; 11-C
plumose, with 9-13 lateral branches. Antenna: Spicules conspicuous; hair 4-A sing-
le, plumose. Thorax: Integument apparently not spiculose; hair 2-P moderately long,
plumose, with numerous lateral branches; 8-P long, shaft not notably thickened,
lateral branches short and moderately numerous; 8-M long, all lateral branches short;
8-T a normal plumose hair; 9-P,M,T barbed. Abdomen: Integument apparently not
spiculose; hair 1-I very short, single or double, 1-II-VII palmate; 6-IV-VI shorter
and less strongly plumose than 6-I-III; hair 9-I-VI distinctly plumose; 13-II-V,VII
rather weakly developed with numerous (4-12) fine branches apically; 4-[V,V weak-
ly plumose; 5-VII long, plumose; 7-VII short, plumose or forked apically; 10-VI
plumose. Segment VIII: Hair 3 plumose; hair 5 4,5f. Anal Segment: Spicules of sad-
dle moderately conspicuous, located along caudal margin and ventral margin distad
of hair 1; hair 1 single, weakly plumose apically.
14 Contrib. Amer. Ent. Inst., vol. 5, no. 2, 1969
SYSTEMATICS. An. fausti is readily separated from all species except arbori-
colus in all known stages by the characters given in the keys. It is distinguished
from arboricolus only in the larval stage; the differences are indicated in the key
and the systematics section of arboricolus.
The distal portion of many of the larval hairs of fausti is plumose or forked;
because of the fineness of these branches it has not been possible to count them
accurately.
In contrast to the other austral species of New World treehole Anopheles, fausti
appears to be a low elevation form.
BIONOMICS. The larvae of the type series were collected in treeholes at eleva-
tions of less than 200 meters in a tropical rain forest. Nothing. is known of the
habits of the adults.
DISTRIBUTION (fig. 1). Known definitely only from Tamazunchale and envi-
rons, at the base of the Sierra Madre Oriental in central Mexico. I have not seen
the material upon which the Nicaragua record in Stone, Knight and Starcke (1959:
18) is based. Material examined: 11 specimens; 2 males, 3 male genitalia, 2 females,
4 larvae.
MEXICO. San Luis Potosi: Tlapexhuacan, near Tamazunchale, Apr 1942, M. Macias, 1 %
[HOPK], 2 6, 1 d gen, 1 9,1 L [ISET], 2 gen, 3 L [USNM].
4. Anopheles (Anopheles) arboricolus Zavortink, n.sp.
‘Figs. 1,8,12
TYPES: Holotype larva (PA 647), “Bajo Grande,” near Cerro Punta, Chiriqui, Panama, eleva-
tion about 2070 meters, from treehole, 18 Mar 1964 [USNM] . Paratypes: 5 larvae (PA 647), same
data as holotype [UCLA].
Anopheles (Anopheles) fausti of Stone, Knight and Starcke (1959:18, in part); Zavortink (1969:
30, in part).
2Anopheles (Anopheles) xelajuensis of Lane (1953:171-172, in part); Stone, Knight and Starcke
(1959:30, in part).
?Anopheles xelajuensis of Galindo (1947:23).
FEMALE. Unknown.
MALE. Wing: 3.67 mm. Proboscis: 2.54 mm. Forefemur: 2.23 mm. Abdomen:
about 2.3 mm. A small dark species. Very similar to fausti and possibly not dis-
tinguishable from it since the following minute differences may be individual rather
than specific. Head: Erect scales short, especially mesally, patch of bright white
scales larger than in fausti. Legs: Femora, tibiae and tarsi dark scaled except for a
ring of semierect white scales at apex of hindfemur, this ring narrower and with
the scales less outstanding than in fausti.
MALE GENITALIA (fig. 12). Probably indistinguishable ca fausti. Sidepiece:
More or less short conical in shape, short; parabasal spines considerably stronger
than internal spine; internal spine more strongly developed than large bristles of
sidepiece, apex recurved. Clasper: Without spicules near base.
PUPA. Unknown.
FOURTH INSTAR LARVA (fig. 8). Head: 0.83 mm. Anal Saddle: 0.35 mm.
Head: Inner clypeals (2-C) closely approximated, separated by a distance much less
than that between inner and outer clypeals, single and simple; outer clypeals (3-C)
distinctly laterad of 4-C, fine, single or forked apically; 11-C plumose, with 6-8 lat-
Zavortink: Treehole Anopheles of New World 15
eral branches. Antenna: Spicules apparently absent; hair 4-A simple, single or dou-
ble. Thorax: Integument apparently without spicules; hair 2-P moderately long, plu-
mose, with numerous lateral branches; 8-P moderately long, shaft very thick, lateral
branches short and numerous; 8-M moderately long, all lateral branches moderately
long; 8-T a normal plumose hair; 9-P,M simple; 9-T very weakly plumose, with 4-6
lateral branches. Abdomen: Integument apparently without spicules; hair 1-I very
short, usually single (1-3b), hair 1-II multiple or palmate, 1-III-VII palmate; 6-I-VI
similar in development and length; 9-I,I] moderately developed, with multiple branch-
es arising near base, 9-III-VI distinctly plumose; 13-II-V,VII moderately developed,
usually 3b (2-6); hair 4-IV,V usually weakly plumose; 5-VII moderately long with
3,4b usually arising near base; 7-VII short with 4-7b usually arising near base; 10-
VII plumose. Segment VIII: Hair 3 single, simple; hair 5 usually with 2,3b from
base. Anal Segment: Spicules on saddle inconspicuous, located along ventral margin
distad of hair 1 and along caudal edge; hair 1 single, simple. |
SYSTEMATICS. An. arboricolus is at present definitely known from a single col-
lection of 3 fourth and 3 third instar larvae from the Chiriqui Volcano region of
western Panama. Although these larvae agree with those of fausti in many signifi-
cant details, they differ in so many others that I do not hesitate to recognize them
as a distinct species in the absence of associated adults. In addition to the characters
given in the key, larvae of the 2 species differ in morphology of hairs 4-A, 8-P,M,
9-P.M,T, 6-IV-VI, 5-VII and 3-VIII. The description of the male and male genitalia
of arboricolus is based on a single individual; the presumptive association of this
specimen with the larvae is probably correct since the specimen is from the same
region as the larvae and, like them, similar to fausti. In fact, as indicated in the des-
cription above, adults and male genitalia of fausti and arboricolus are probably in-
distinguishable.
The original record of xelajuensis from the Chiriqui Volcano region of Panama
(Galindo, 1947:23) and the subsequent ones based on it (Lane, 1953:172; Stone,
Knight and Starcke, 1959:30) possibly refer to arboricolus. The record of fausti
from Panama (Stone, Knight and Starcke, 1959:18) is apparently based on the same
male that I have seen and tentatively associated with arboricolus.
BIONOMICS. The larvae of the type series were collected in a treehole at an ele-
vation of about 2070 meters. The report of finding 6 males and 4 females of xela-
juensis resting in hollow trees at an elevation of about 1900 meters on Volcan Chiri-
qui (Galindo, 1947:23) may refer to this species.
DISTRIBUTION (fig. 1). Currently known only from high elevations in the Chiri-
qui Volcano area of western Panama. Material examined: 8 specimens; 1 male, 1
male genitalia, 6 larvae.
PANAMA. Chiriqui: “Bajo Grande,” near Cerro Punta, type series, see above. El Volcan Chiri-
qui, 30 June 1943, T.H.G. Aitken, 1 3, 1 é gen [USNM].
5. Anopheles (Anopheles) xelajuensis de Leon
Pigs. 1 910,12
1938. Anopheles xelajuensis de Leon, 1938:421-423. TYPE: Holotype 6, Cerro Quemado, near
Quezaltenango, Quezaltenango, Guatemala, 2 Jan 1936, J. Romeo de Leon [ESPG].
Anopheles (Anopheles) xelajuensis of Lane (1953:171-172, in part); Stone, Knight and Starcke
(1959:30, in part); Belkin, Schick and Heinemann (1965:28); Zavortink (1969:30).
16 Contrib. Amer. Ent. Inst., vol. 5, no. 2, 1969
Anopheles xelajuensis of Komp (1941:92,96); Russel, Rozeboom and Stone (1943:35, in part);
Forattini (1961:174,180,181).
Anopheles (Coelodiazesis) xelajuensis of Vargas and Martinez (1956:116-119,142); Vargas (1959:
377).
Anopheles (Russellia) xelajuensis of Vargas (1943:65 ,67 ,68-70); Senevet (1958:44).
FEMALE (fig. 12). Wing: 5.16 mm. Proboscis: 2.92 mm. Forefemur: 2.75 mm.
Abdomen: about 2.6 mm. A large dark species. Head: Integument dark brown; in-
terocular bristles whitish; erect scales long, mostly black, anterior ones brilliant
white; interocular scales white, much elongate, forming a conspicuous tuft; palpus
entirely dark scaled, basal scales semierect. Thorax: Mesonotum long and only slight-
ly arched; mesonotal integument dark brown with a broad hoary median longitudi-
nal stripe; pleural integument dark brown; anterior mesonotal bristles not as long,
strong or conspicuous as in barberi and judithae, dark except for light anterior acros-
tichals; narrow white scales in tuft in center of anterior promontory and extending
through anterior third of acrostichal line; ppl bristles 3-5. Legs: Coxae and trochan-
ters white; anterior forecoxal bristles 8-16; femora, tibiae and tarsi dark scaled ex-
cept as follows: femora with apical white scales, few on foreleg, many, forming a
conspicuous ring, on hindleg; fore- and midtibiae with a few white scales at apex;
hindtibia with long broad apical white patch covering anterior, dorsal and posterior
surfaces. Wing: Largely dark scaled; apex of costa and vein R, with yellowish-white
scales; 4 fringe spots usually present, 1 large, extending from end of costa to vein
R45, 3 smaller, at ends of veins M,4,, M344 and Cu,; dark scales not uniformly
distributed, but grouped into strong spots at base of veins Rs and R4+s5 and at furca-
tion of veins R,43, M and Cu; profusely scaled; scales in middle portion of vein 1A
somewhat spreading. Abdomen: Tergites black, sternites black with basal white
band.
MALE. As for female except for sexual characters.
MALE GENITALIA (fig. 9). Sidepiece: More or less cylindrical in shape, long;
spicules arranged singly, in rows or in small clumps; scales numerous, 6-16; para-
basal spines stronger than internal spine; lateral parabasal spine not flattened pre-
apically, apex gradually attenuate and recurved; internal spine slightly stronger than
large bristles of sidepiece, apex curved. Claspette: Lateral 2 setae of ventral lobe
subequal in stoutness, but outermost usually shorter. Clasper: With at least a few
spicules near base of ventral surface.
PUPA (fig. 9). Abdomen: not measured. Trumpet: 0.55 mm. Paddle: 1.19 mm.
Width of segment VIII: 0.92 mm. Cephalothorax: Moderately pigmented with dark-
er areas more extensive than in powderi; hair 3-C much longer than 1-C. Trumpet:
Brown. Abdomen: Moderately pigmented, darker areas more extensive than in pow-
deri; hair 6-I much longer than 7-I; hair 1-II,III] multiple (3-5b); hair 5-III,IV short,
fine, usually 3,4b; hair 5-V-VII thickened, much longer than hair 9 of correspond-
ing segment, and with 3-5 fine lateral or apical branches; 9-III-VIII moderately pig-
mented, concolorous with to slightly darker than abdominal integument; 9-III ap-
proaching 9-IV in both diameter and length; 0-VII enlarged, 5,6b; hair 7-VII single,
longer than 9-VIII. Paddle: Moderately pigmented, midrib not darker.
FOURTH INSTAR LARVA (fig. 10). Head: 0.90 mm. Anal Saddle: 0.39 mm.
Head: Inner clypeals (2-C) closely approximated, separated by a distance much less
than that between inner and outer clypeals, single and simple; outer clypeals (3-C)
distinctly laterad of 4-C, single, thickened; 11-C not plumose, with 2-5b from near
base. Antenna: Spicules inconspicuous; hair 4-A single and simple. Thorax: Integu-
Zavortink: Treehole Anopheles of New World M7
ment, at least on ventral surface, spiculose; hair 2-P stellate, 9-15b; hair 8-P moder-
ately long, shaft very stout, lateral branches very short and moderately numerous;
8-M moderately long, basal lateral branches very long; 8-T with thickened shaft, not
plumose to apex, distal end blunt; 9-P,M,T barbed. Abdomen: Integument, at least
on more posterior segments, spiculose; 1-I-VII stellate; 6-IV-VI less strongly plumose
than 6-I-III; hair 9-I-VI stellate, very strongly developed; 13-II-V,VII strongly devel-
oped, stellate; 4-IV,V simple, single or double; 5,7-VII stellate, 8-11b; hair 10-VII
stellate, 4-6b. Segment VIIT: Hair 3 with 3-6 very stout branches from base; hair 5
stellate, 18-22b. Anal Segment: Spicules on saddle very conspicuous, located on all
but basal portion of saddle; hair 1 with 3-5b, usually from base.
SYSTEMATICS. An. xelajuensis is the most differentiated New World treehole
Anopheles. The most striking characteristics of the species have been indicated in
the keys and the systematics chapter.
According to the original description, which is based on a single male from Guate-
mala, the hindtibia of xelajuensis is entirely dark scaled. The adults from Mexico
examined during this study have a conspicuous large white patch at the apex of the
hindtibia. It is possible that these specimens represent a different species, but this
cannot be determined until additional material, especially topotypic xelajuensis, is
obtained. :
BIONOMICS. Larvae of xelajuensis have been collected in a treehole at an eleva-
tion of 2500 meters. According to Vargas (1943:70) the larvae feed on bottom sedi-
ments and spend relatively little time at the surface, the pupal stage lasts 12 to 15
days and females readily bite humans. The holotype male was collected between
rocks in an oak forest in the highlands of Guatemala.
_ DISTRIBUTION (fig. 1). Known only from high elevations in the Sierra Madre
del Sur of southern Mexico and the Sierra Madre in Guatemala. The Panama rec-
ords of Galindo (1947:23), Lane (1953:172) and Stone, Knight and Starcke (1959:
30) refer to arboricolus or powderi. Material examined: 13 specimens; 2 males, 3
male genitalia, 1 female, 1 pupa, 6 larvae.
MEXICO. Oaxaca: Galera Vieja, between Ixtlan and Tepanzacoalco, Sept 1942, M. Macias, 1 6
gen, 1 L [CU],24,2¢ gen, 19,2 L [ISET], 1 P, 2 L [USNM], 1 L [UCLA].
6. Anopheles (Anopheles) powderi Zavortink, n.sp.
Figs. 1,11
TYPE: Holotype ¢ with associated pupal skin (CR 50-102), about 10 km SE of La Sierra, San
Jose, Costa Rica, elevation about 2400 meters, from rothole in fallen tree, 24 Nov 1962, C.L.
Hogue and W.A. Powder [USNM]. This species is dedicated to William A. Powder, in recognition
of his contributions to the “Mosquitoes of Middle America” project.
?Anopheles (Anopheles) xelajuensis of Lane (1953:171-172, in part); Stone, Knight and Starcke
(1959:30, in part).
?Anopheles xelajuensis of Galindo (1947:23).
FEMALE (fig. 11). Wing: 5.15 mm. Proboscis: 3.13 mm. Forefemur: 2.88 mm.
Abdomen: about 2.7 mm. A large dark species. Quite similar to xelajuensis, differ-
ing mostly in ornamentation of leg and wing. Head: Interocular bristles amber. Tho-
rax: Whitish acrostichal scales extending about halfway to scutellum. Legs: Femora,
tibiae and tarsi dark scaled except for conspicuous ring of semierect white scales at
18 Contrib. Amer. Ent. Inst., vol. 5, no. 2, 1969
apex of hindfemur and a few white scales at apex of hindtibia. Wing: Veins and
fringe dark scaled except for a single large apical fringe spot extending from end of
vein R, to end of vein M,+4,; scales of veins uniformly distributed and not grouped
into dark spots; profusely scaled; scales in middle portion of vein 1A wide spread-
ing. Abdomen: Tergites and sternites black.
MALE and MALE GENITALIA. Unknown.
PUPA (fig. 11). Abdomen: 4.75 mm. Trumpet: 0.54 mm. Paddle: 1.27 mm.
Width of segment VIII: 1.11 mm. Cephalothorax: Lightly pigmented with darker
areas less extensive than in xelajuensis; hairs 1,3-C subequal in length. Trumpet:
Light amber in color. Abdomen: Lightly pigmented, darker areas less extensive than
in xelajuensis; hairs 6,7-I subequal in length; 1-II,III multiple; 5-III-VII short, fine,
usually 2,3b; hair 9-III-VIII lightly pigmented, more or less concolorous with ab-
dominal integument; 9-III approaching 9-IV in diameter and length; O-VII small,
single; 7-VII double, shorter than 9-VIII. Paddle: Very lightly pigmented, midrib
not noticeably darker.
LARVA. Unknown.
SYSTEMATICS. An. powderi is known from a single female with its associated
pupal skin. The adult, as indicated in the key, is adequately distinct but the pupa,
except for its much larger size, is scarcely differentiated from that of judithae. An.
xelajuensis is the only other New World treehole Anopheles of comparable size.
This species could extend through the Cordillera de Talamanca into western Pan-
ama, and, if so, it might be the xelajuensis of Galindo (1947:23), Lane (1953:172)
and Stone, Knight and Starcke (1959:30).
BIONOMICS. The pupa of powderi was taken from a rothole in a fallen tree at an
elevation of about 2400 meters in a cloud forest.
DISTRIBUTION (fig. 1). Presently known solely from the holotype which was
collected high in the Cordillera de Talamanca in southern Costa Rica. Material ex-
amined: 2 specimens; 1 female, 1 pupa; 1 pupal rearing.
COSTA RICA. San Jose: La Sierra (10 km SE), holotype, see above.
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Belkin, John N., R.X. Schick and S.J. Heinemann
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Howard, Leland O., H.G. Dyar and F. Knab
1917. The mosquitoes of North and Central America and the West Indies. v.4.
Wash., Carnegie Inst. (Publication 159). p. 525-1064.
Jenkins, Dale W. and S.J. Carpenter
1946. Ecology of the tree hole breeding mosquitoes of Nearctic North America.
Ecol. Monogr. 16:31-47.
Komp, William H.W.
1941. The classification and identification of the Anopheles mosquitoes of Mexi-
co, Central America and the West Indies. Jn Moulton, Forest, R., A symposium
on human malaria. Wash., Amer. Ass. Advance. Sci. (Publication 15). 398 p.
Lane, John
1953. Neotropical Culicidae. Sao Paulo, Univ. Sao Paulo. 2 v. 1112 p.
de Leon, J. Romeo
1938. El anophelismo de altura en Guatemala. Bol. ai Gen. Sanid. Publica (Gua-
temala) 9:411-424.
Matheson, Robert
1944. Handbook of the mosquitoes of North America. ed. 2. Ithaca, Comstock
Publishing Co. 314 p.
Nielsen, Lewis T., J H. Arnell and J.H. Linam
1967. A report on the distribution and biology of the tree hole mosquitoes in the
western United States. Calif. Mosquito Control Ass., Proc. Pap. 35:72-76.
Nielsen, Lewis T., JH. Linam, J H. Arnell and T.J. Zavortink
1968. Distributional and biological notes on the tree hole mosquitoes of the west-
ern United States. Mosquito News 28:361-365.
Penn, George H.
1949. Pupae of the Nearctic anopheline mosquitoes north of Mexico. Nat. Malar-
ia Soc., J. 8:50-69.
20 Contrib. Amer. Ent. Inst., vol. 5, no. 2, 1969
Petersen, J.J., H.C. Chapman and O.R. Willis
1969. Predation of Anopheles barberi Coquillett on first instar mosquito larvae.
Mosquito News 29:134-135.
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. Mosquito News 16:10-17.
Ribgy, Paul T., T.E. Blakeslee and C.E. Forehand
1963. The occurrence of Aedes taeniorhynchus (Wiedemann), Anopheles barberi
(Coquillett), and Culex thriambus (Dyar) in Arizona. Mosquito News 23:50.
Russel, Paul F., L.E. Rozeboom and A. Stone
1943. Keys to the anopheline mosquitoes of the World. Philadelphia, Amer. Ent-
omol. Soc. 152 p.
Senevet, Georges
1958. Les Anopheles due globe. Revision generale. Encycl. Entomol. (A) 36.
215 p:
Stone, Alan, K.L. Knight and H. Starcke
1959. A synoptic catalog of the mosquitoes of the World (Diptera, Culicidae).
Wash., Entomol. Soc. Amer. (Thomas Say Found. Publication 6). 358 p.
Stratman-Thomas, W.K. and F.C. Baker
1936. Anopheles barberi Coquillett, as a vector of Plasmodium vivax Grassi and
Feletti. Amer. J. Hyg. 24:182-183.
Vargas, Luis
1940. Anopheles (Anopheles) barberi en Mexico. Inst. Salubr. Enferm. Trop.,
Rev. $:319-322..,.
1942a. Anopheles xelajuensis Romeo de Leon, 1938 en Mexico. Inst. Salubr. En-
ferm. Trop., Rev. 3:169-175.
1942b. El huevo de Anopheles barberi Coquillett, 1903. Inst. Salubr. Enferm.
Trop., Rev. 3:329-331.
1943. Los subgeneros Americanos de Anopheles (Diptera, Culicidae). Anopheles
(Russellia) xelajuensis de Leon, 1938 n.subgn. y Anopheles (Coelodiazesis)
fausti n.sp. Inst. Salubr. Enferm. Trop., Rev. 4:57-77.
1959. Lista de Anopheles de las Americas y su identificacion por caracteres mas-
culinos. Inst. Salubr. Enferm. Trop., Rev. 19:367-386.
Vargas, Luis and Amado Martinez Palacios
1956. Anofelinos Mexicanos Taxonomia y distribucion. Mexico, D.F., Secretar.
Salubr. Asistencia. 181 p.
Zavortink, Thomas J.
1969. Mosquito Studies (Diptera, Culicidae). XV. A new species of treehole breed-
Anopheles from the southwestern United States. Amer. Entomol. Inst., Con-
trib. 4(4):27-38.
OOAYANNBWNHN
aa
Zavortink: Treehole Anopheles of New World pA |
FIGURES
Distribution of collections examined
Anopheles (An.) barberi; adult
Anopheles (An.) barberi; male genitalia and pupa
Anopheles (An.) barberi; larva
Anopheles (An.) judithae; male genitalia and pupa
Anopheles (An.) judithae; larva
Anopheles (An.) fausti; larva
Anopheles (An.) arboricolus; larva
Anopheles (An.) xelajuensis; male genitalia and pupa
Anopheles (An. ) xelajuensis; larva
. Anopheles (An.) powderi; pupa and pleuron and mesonotum of adult
. Anopheles (An.) arboricolus; male genitalia. Anopheles (An.) fausti; male gen-
talia, wing and hindleg. Anopheles (An.) xelajuensis; wing and hindleg.
SATIN ALAIV.LS
00s
Oot os 0
oor ooros 0
SHILAWO TA (Rx.
WAN WIGGIN +
we
¢ a
Te
+Ssisuan{Djex v
llapmMOd w
apypipnt O
SND ©
l4aqudg @
snjoodli0qin @
SSTSHAON
@ ANOPHELES <==
TAM =< —
WAAR SS =
SES =
Se =
A'S —s
DT Win 2 ns . Pir te ape \ RYNANAWN
i; PAHO ALAINMATTATRMANK GU Nw
HAIN it it a ll ut Ls \\
HUME, if PEmeN nee
x
Wi
WMA
ie Mi
1
Mf} VY 4
:
SAN EOS
Z Hae WN
barberi *@ y)
FORE MID HIND
Maryland
United States
ANOPHELES
ANOPHELES
barberi
UCLA 437
Ohio
United States
ANOPHELES
on
rig SD pupal
Ve WY
ANOPHELES
FF2
HILLY,
WY
7
a
TT
BA
"
fausti
San Luis Potosi
Mexico
ANOPHELES
ANOPHELES
IV
; 1
ma .
= 7 SX
[= ee {14
arboricolus
Chiriqui
Panama
) ANOPHELES
aN
Fig. lO , ANOPHELES
1
10
INGUIN WY
oN WONKA UY
INA IN WY
AN
a
=
AE BE,
Y,
A hoagh phage
°
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S AN
. e
ks =
wo
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A "Zeal
a
aii
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~
rs)
MEA
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San a
LM an
a
<y
hy
ws ae
SWANN ER
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Ny S \ an =a
S AY LSS
|
NSS
= N .
\
Hi
SAIN
Y Es
ue ee: SM
PEE 12
A
a Yc
a
HH
xelajuensis
Oaxaca
Mexico
ANOPHELES
ANOPHELES
SSSTSN
Oaxaca
Mexico
i it nt i
ili Mi
aa
wiuit
\
nen ni ji
iy
NN
2A Le
fausti Jbin ar
aad
WSS
SSSS
San Luis Potosi
Mexico
Mow VY
Wi fi ~ :
WAN \ Ss
\W \\
Ha AY
mt w
Nia i i Nii) Fini am inl iN i co
NAW Nh iY NM
a TT i AN
in ANNA Au
arboricolus
Chiriqui
Panama
fausti
San Luis Potosi
Mexico
=
iN)
TABLE 1.
Groups of New World treehole Anopheles
CHARACTER barberi and judithae arboricolus and fausti
small large
SIZE large
ADULT
interocular scales
absent to numerous numerous and long numerous and long numerous and long
and short
erect head scales light mesally, light or bright white mesally, bright white mesally, bright white mesally,
darker laterally very dark laterally very dark laterally very dark laterally
light palpal patches absent present absent absent
mesonotum shortened and arched longer and flatter longer and flatter longer and flatter
mesonotal hoary absent present present present
stripe
absent or tuft on
promontory
acrostichal scaling long line long line long line
color of coxae same as adjacent por- much lighter than much lighter than much lighter than
tions of pleuron pleuron pleuron pleuron
apex of hindfemur dark conspicuous white conspicuous white conspicuous white
ring ring ring
apex of hindtibia dark dark conspicuous white dark
patch
dark spots on absent absent present at base of absent
wing veins branches
light patches on absent absent present at apex of absent
wing veins costa and R,
light fringe spots absent 1, from Ry to R4a+s5 4,1 from R, to Rays, | 1, from R,; to M442
1 each at end of
Mj +2,M34q and Cu,
MALE GENITALIA
sidepiece
short unknown
large clumps
short
singly, rows or
small clumps
long
arrangement of side-
piece spicules
singly, rows or
small clumps
} unknown
sidepiece scales absent few many | unknown
ventral claspette lobe | lateral seta not as strongly } unknown
developed as 1 next
mesad
lateral 2 setae subequally
developed
lateral seta not as strongly
developed as 1 next
mesad
LARVA
hair 3-C fine, mesad of 4-C fine, laterad of 4-C thick, laterad of 4-C unknown
11 1-3b from base plumose 2-5b from base unknown
1-III-VII palmate palmate stellate unknown
9-III-VI branched from base, mod- unknown
erately developed
plumose stellate, very strongly
developed
4.IV,V
10-VII
simple, single unknown
2,3b from base or middle
plumose simple, single or double
4-6b, stellate
plumose unknown
DISTRIBUTION restricted restricted
southern Mexico,
Guatemala
restricted
widespread
United States, northern
Mexico
central Mexico, west- Costa Rica
ern Panama
HABITAT
montane cloud
forest
temperate deciduous forest, wet montane forest
xeric evergreen forest,
gallery forest
humid montane forest,
tropical forest
Zavortink: Treehole Anopheles of New World
INDEX TO SCIENTIFIC NAMES
arboricolus Zavortink, 2, 3, 5-7k, 14, 14-15, 17; 17, 8, 12
barberi Coquillett, 1, 2, 3, 5-7k, 7-10, 10, 11, 13, 16; 1-4
barberi of authors, 10
barianensis James, 2
Coelodiazesis Dyar & Knab, 2
Cyclophorus Eysell, 2
eiseni Coquillett, 3, 5-6k
fausti Vargas, 1, 2, 3, 5-7k, 12-14, 14, 15; J, 7, 12
fausti of authors, 14, 15 |
judithae Zavortink, 2, 5-7k, 8,9, 10-12, 13, 16, 18; 1, 5, 6
plumbeus Stephens, 2
powderi Zavortink, 2, 3, 5-6k, 16, 17, 17-18; /, 11
Russellia Vargas, 2
xelajuensis de Leon, 1, 2, 3, 4, 5-6k, 9, 15-17, 17, 18; 1, 9, 10, 12
xelajuensis of authors, 13, 14, 15, 17, 18
Bs 5)
(Continued from inside front cover)
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By Robert X. Schick.
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(Continued on back cover)
- MOSQUITO STUDIES (Diptera, Culicidae)
XX. THE TERRENS GROUP OF AEDES (FINLAYA)’
by
Robert X. Schick’?
CONTENTS
Oh TID THE ae Fil GJ Sie eS SOG MON Nae eo My car Pen re CORO amen MGT NENT Bic Ne Pee WP Vata ha INR
Ee Oi GI GH ied PE a mT ca ore re Mee eee myn een iia er SEMEN Sets eOMUPI EN OE
Be NO A TO ii ie i A oe he Seen Ak sk BME cd Bada Ue
ISTO TET Bs Gy CU Gee BSH | 2 SRN PN eee ey ene Oh atone a ene Comme et Ma
ee AACS AND Te LOIN oo oie ok were Meio ai td he Na ace te a
ESOP EN OG LOT RIE MEAs Sie kG MONG SA en A eae ieee rer aaat vile Meine) eoray by mea gas freee
BIONOMICS...... ee ee UA Aenea a a eM GL a eta a agar aca
DISEASE RELATIONS ee A el EUR RON MAY eiacg ay Meet KO ool Watt RTE pay a commen AES)
TAXONOMIC TREATMENT . a ee nd au Sal) eal SEN ama alae whic nace
“USEC GTO) CARAS GB SAT ee) Gerace ATE Ey Ue na LCR ON IRN Ry REIL 1
Te EO SOCIO ee ai i ay i Nek a I ae a me br ak alan ne ae am
Thorntoni Subgroup. . TTT AT GREE ai tot es 0 SO gana Ni Hea Ep)
1. Aedes (F.) thorntoni Dyar & Knab . ae MO IGE ONG Re ate
2. Aedes (F.) argyrothorax PEE Ey cha
Bertrami Subgroup . . . Le ae PRR a ea ae.
3. Aedes (F.) bertrami, n. sp. SL NEE NIE SEEMS IR Scie nS RN Pat REM NIGES
Terrens Subgroup . . Pa a Aiea a ee Lat ata aa SSL gt a ada ee
4. Aedes (F.) terrens (aiken nee Se ae Re TAL ga SL a Deans ear eat =|
5. Aedes (F.) braziliensis Gordon & Poy
ON ACO CRIT D7 AV ROR IR ig i ee eae ae nae uu eo
es ICS OUI UID i a igen ok wel Wiha a ala Lue th gies aaa ae
Be CO Mi ROMO I el) lakes Lae oa Male ky ue ea a ack ee
Alboapicus Subgroup .. . pits Ea NRT te a aR ad anes
9. Aedes (F.) alboapicus, n. sp. BE OR AP aS IRR aI St ee Bo
‘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. 5, no. 3, 1970
Buenaventura Subgroup . :
10. Aedes (F.) buenaventura, n. sp.
Metoecopus Subgroup .
11. Aedes (F.) metoecopus Dyar.
Insolitus Subgroup . .
12. Aedes (F.) ae (Coquillett
Aitkeni Subgroup...
13. Aedes (F) aitkeni, n. sp.
Homoeopus Subgroup . :
14. Aedes (F.) homoeopus Dyar .
15. Aedes (F.) impostor, n.sp.
Heteropus Subgroup
16. Aedes (F.) amabilis, n. Sp.
17. Aedes (F.) gabriel, n.sp.
18. Aedes (F.) idanus, n.sp.
19. Aedes (F.) sumidero, n.sp.
20. Aedes (F.) vargasi, n.sp.
21. Aedes (F.) heteropus a
Galindoi Subgroup . . .
22. Aedes (F.) galindoi, n. AD.
23. Aedes (F.) campana, n.sp.
24. Aedes (F.) daryi, n.sp. .
Podographicus Subgroup. . .
25. Aedes (F.) ea. Dyar & Knab
Tehuantepec Subgroup
26. Aedes (F.) tehuantepec, n. sp.
27. Aedes (F.) schroederi, n.sp.
Diazi Subgroup. . ;
28. Aedes (F.) diazi, n. sp.
REFERENCES CITED
FIGURES
INDEX TO SCIENTIFIC 1 NAMES
Schick: Terrens Group of Aedes (Finlaya) 3
INTRODUCTION
The Terrens Group of the subgenus Finlaya of Aedes is Neotropical in distribu-
tion and forms the dominant group of Finlaya of the region. Twenty-eight species
are recognized in the present revision, as opposed to only 2, terrens and argyrothor-
ax, in the world catalog (Stone, Knight and Starcke, 1959). The adults can usually
be recognized by the characteristic silver banding of the tarsi, tarsi 1-II,[JI showing
basal and apical silver bands and tarsi 2-II,III only a basal silver band. The pupae do
not show distinctive group characters and the larvae are distinguished only by a com-
bination of several characters which are given in the description of the group. All
the species are forest dwellers and utilize container habitats for breeding, primarily
treeholes.
The present revision is based on a study of 3561 specimens, 946 9, 772 6,945 pu-
pae, 1097 larvae, with 542 individual rearings (329 larval, 178 pupal, 35 incom-
plete). Most of this material was collected for the project ‘““Mosquitoes of Middle
America” (Belkin, Schick et al, 1965) utilizing the techniques outlined by Belkin,
Hogue et al (1965) and is deposited in the Department of Zoology, University of
California, Los Angeles (UCLA). The remaining material was obtained from the U.S.
National Museum (USNM) where the types of all the new species will be deposited.
The methods of presentation in the present paper (format, synonymies, descriptions,
distributional data and illustrations) are generally similar to those used by Berlin
(1969:4-5). Modal ranges, that is 75% of the values about the median, are given for
the various mensurations in addition to the absolute ranges, the absolute range be-
ing indicated parenthetically following the modal value. An effort was made to cite
the elevation of all localities in the distributional data. When not specifically given
in the collection data the elevations were determined mainly from contour maps
and are enclosed by brackets. Usually only the range between contour lines could be
obtained; these elevations were rounded to the nearest 100 feet.
The mesonotum of both sexes, male genitalia and immature stages are routinely
illustrated for each species. These structures and stages are also illustrated for differ-
ent geographical populations of a few possibly polytypic species. The illustrations
of the mesonota show the pattern and different intensities of silver scalation dia-
grammatically by the use of screens of 3 different low densities; dark scaled or bare
areas are represented by a very dense screen. All illustrations are based upon speci-
mens from the type locality or from as near the type locality as possible. The pupal
and larval drawings show a modal condition determined respectively from at least 5
and 3 specimens usually of the same collection. The modal number of branches
and usual length of those hairs treated in the descriptions were based upon all avail-
able specimens or upon a representative sample from each collection.
Measurements were made with the aid of American Optical net reticule 1409A
ruled into squares of 0.5 mm and were usually determined from all available speci-
mens. Only reared specimens were utilized in those cases in which there was an
abundance of material in a particular collection or from a particular geographic area.
The terminology in general follows Belkin (1962) except that a few special terms
are introduced for the pattern of adult ornamentation, some of the structural spec-
ializations of the male genitalia and several indices and ratios. These are explained
in the section on Taxonomic Characters.
I am indebted to John N. Belkin for providing descriptive notes on the type speci-
mens in the British Museum (Nat. Hist.) and the U.S. National Museum of the fol-
lowing species: braziliensis, heteropus, homoeopus, insolitus, metoecopus, podo-
& Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
graphicus, terrens and thorntoni, and also for advice. I thank Thomas J. Zavortink
and O.G.W. Berlin for their assistance; William A. Powder and Sandra J. Heinemann
for the preparations of the immature stages and male genitalia; Rainer Beck, Sharon
Burmeister, Sally Dieckmann, Nobuko Kitamura, Margaret Kowalczyk and Nancy
Martsch for the preliminary and/or final illustrations; Sheila Bernstein for some of
the preliminary drafts of the manuscript and Caryle Abrams for other preliminary
drafts and the final copy for reproduction. I also thank T.H.G. Aitken, Pedro
Galindo, Charles L. Hogue, Vernon Lee and Charles Serie, who made possible
through their cooperation many of the collections on which this study is based;
and Alan Stone for making available the material in the U.S. National Museum.
KOMP COLLECTION
The Komp collection, part of which is housed at the USNM and part at UCLA,
requires some explanation in regard to the numbering system. Most of the specimens
were collected by Komp himself, primarily in the Canal Zone in the years 1943 and
1945, and the remaining specimens were collected by other workers in Central
America and northern South America. Collection data (when known) accompany
all the specimens, both pinned and on slides. The adults and skins of the reared pu-
pae and larvae were assigned numbers and these numbers and the collection data
were typed on 3x5 index cards (usually in duplicate). Komp used 2 different num-
bering systems, 1 for the rearings and 1 for the adult specimens, reared or not. Un-
fortunately Komp did not give cross references on the specimens so that the cards
are necessary to locate associated. stages. The numbers for the rearings of the year
1943 are in chronological order; these numbers are noted in 3 different ways, e.g.
for rearing 92, No. 92, No. 3-92 or No. 43-92. A capital letter A, P or L follows
some of the numbers and apparently refers to the specific stage. The few rearings
for the year 1944 have a “44” prefix and all the 1945 rearings a “5” prefix (e.g.
No. 5-163). Unlike the 1943 system the 1945 rearing numbers are not arranged in
any particular order, although some of the earlier rearings tend to have lower num-
bers. Most of the rearing numbers represent single individual rearings but some rep-
resent 2 or 3. In the latter cases, all the skins are mounted on the same slide; some-
times 1 or more of the pupal skins are absent and sometimes more than | species
is represented.
All the adults of the Terrens Group were given “‘serial’? number 207 and there
are 5 sets of these numbers, 207A,B,C,D and E. A complete set basically comprised
45 adult numbers, 207A-1, 207A-2...207A-45; 207B-1-45 and so on. Adult num-
bers 207A and B represent collections made in 1943 and earlier, and includes most
of Komp’s 1943 collections and his 3 1944 collections; 207C represents collections
made by Komp in 1945; and 207D and E collections made at various times and in-
cludes some of his 1943 and many of his 1945 collections. Komp, when assigning
the adult numbers, selected the adults either at random or in some order not obvi-
ous to me. The adult numbers represent either reared or wild caught adults; in the
latter case several adults may be represented by a single number. Only the adult
numbers are used in the present revision in the distributional lists.
Although Komp’s rearing numbers usually represent only a portion of a given col-
lection several series of rearings can be assigned to a particular collection by using
Komp’s data. These data are the date of the collection and the specific treehole
from which the specimen was collected (e.g. “‘treehole No. 1 SM [Snyder Molino]
Schick: Terrens Group of Aedes (Finlaya) 5
trail” or “small treehole opp. WMW [William M. Wheeler trail] 14’’). Thus, in the
distribution sections of thorntoni, zavortinki, alboapicus and galindoi a single col-
lection may be represented by several apparently unrelated adult numbers.
TAXONOMIC HISTORY
Aedes terrens was described by Walker in 1856 as a Culex but the true identity of
the species was not known until 1921. The second species, oswaldi, was described
by Lutz in 1904 as a Gualteria. With the exception of Dyar and Knab (1906a:166),
who placed oswaldi in Haemagogus, Gualteria was retained until Howard, Dyar and
Knab (1917:815) transferred oswaldi to Aedes. Coquillett in 1906 described jnsolitus
and Jaternaria in the genus Verrallina but these species were placed in Aedes in the
same year by Dyar (1906:16). All the other species of the group were originally des-
cribed in Aedes. Dyar (1917:79), on the basis of the male genitalia, placed all the
species of the Terrens Group known at that time in the subgenus Gualteria of Aedes
(together with A. triseriatus and A. mediovittatus). These were subsequently trans-
ferred to the subgenus Finlaya by the same author in his Mosquitoes of the Amer-
icas (Dyar, 1928). Edwards (1932) placed the Terrens Group species into Finlaya
Group B (terrens-gubernatoris-group: Gualteria). Knight and Marks (1952) revised
and expanded this scheme. Group B was referred to as TERRENS-group: GUAL-
TERIA and was subdivided into several subgroups of which A and B comprise the
Terrens Group of the present revision. Vargas (1950) resurrected the subgenus Gual-
teria for the Terrens Group and other species, both of the New and Old World,
based upon differences between these species and poicilius (Theobald, 1903), which
he considered as the type species of Finlaya. This change has been accepted by a
few South American workers but Finlaya remains the generally accepted subgeneric
name for the group.
The taxonomy of the species, based principally upon the adults, has been in
a state of confusion since the first general review in 1917 by Howard, Dyar and
Knab. Five nominal species were known at that time: oswaldi Lutz, 1904; inso-
litus (Coquillett, 1906); podographicus Dyar & Knab, 1906; laternarius (Coquil-
lett, 1906); and thorntoni Dyar & Knab, 1907. Three valid species were recognized
by Howard, Dyar and Knab and were characterized principally upon the basis of
the presence or absence of transverse silvering on the mesonotal disc: (1) thorntoni,
mesonotal disc transversely silvered in both sexes, (2) oswaldi, disc transversely sil-
vered only in the male and (3) podographicus, disc not transversely silvered in either
sex. The Coquillett species were incorrectly synonymized with oswaldi and this laid
the foundation for much of the confusion in subsequent works. The oswaldi of
Howard, Dyar and Knab was essentially based upon insolitus and true oswaldi would
key to podographicus in their work. The mesonotal disc in oswaldi is actually not
transversely silvered in either sex while the disc in insolitus (=laternarius) is trans-
versely silvered only in the male.
Dyar in 1921 published a 3 page revision of the group, the only one ever devoted
solely to the Terrens Group. The 1921 revision differed from the one of 1917 in 2
respects: (1) two species were added, heteropus, a new species, showing the mes-
onotal scalation of the podographicus type (and correctly distinguished from that
species by the broader median dark band of tarsus 1-II), and argyrothorax Bonne-
Wepster & Bonne, 1919, known only by the male, showing a transversely silvered
6 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
disc and a characteristic claspette filament; and (2) the name terrens was used for
oswaldi. The true identity of Aedes terrens had been discovered by Bonne-Wepster
and Bonne (1921:23) and these authors considered terrens related to but distinct
from oswaldi. Dyar correctly synonymized the 2, and insolitus and laternarius thus
also became synonyms of terrens.
Three more species were described between 1921 and 1928, the date of the next
revision, terrens homoeopus Dyar, 1922; terrens var. braziliensis Gordon & Evans,
1922; and metoecopus Dyar, 1925. The first 2 species were considered as forms of
the terrens of Dyar since the mesonotal disc of the male was transversely silvered
and the claspette filament was not of the argyrothorax type. Aedes homoeopus was
distinguished from terrens on the basis of valid characters of the male genitalia and
braziliensis on the basis of various characters of apparently no taxonomic value, al-
though this species is actually distinct. Aedes metoecopus showed the mesonotal
ornamentation of the podographicus and heteropus type and was distinguished
from these species on the basis of valid characters of the male genitalia.
Dyar (1928:223-226) showed less insight into the group than in his previous pub-
lications. Only 4 valid species were recognized: argyrothorax, thorntoni, terrens,
and podographicus. These were characterized as in the 1917 and 1921 publications.
Aedes homoeopus and braziliensis were added to the list of terrens synonyms and
heteropus and metoecopus were synonymized with podographicus. The reasons for
these incorrect synonymies were not discussed. As in the case of the 1917 revision,
true terrens keys to podographicus and this explains its identification as podograph-
icus by Costa Lima (1930:257) and Lane (1936a:11; 1936b:131).
The treatment of the species by Edwards (1932) in the Genera Insectorum served
only to confuse the picture further. Three valid species were recognized, argyrotho-
rax, thorntoni and terrens. Aedes terrens comprised 3 varieties, terrens, metoecopus
and podographicus; the variety terrens included oswaldi and braziliensis, and the var-
iety podographicus included insolitus, laternarius, heteropus and homoeopus. | can
see no justification in the changes made by Edwards and no justification was given.
The variety podographicus in the sense of Edwards is quite heterogeneous compared
to the other taxa recognized by him.
Lane (1939:105) and Kumm, Komp and Ruiz (1940:416) took the ultimate step
in the trend of synonymizing valid species and lumped all the species, except argyro-
thorax, under terrens. The latter authors stated “‘we agree with Shannon and Ed-
wards that thorntoni, podographicus and terrens [the latter 2 species apparently in
the broad sense of Dyar (1928)] are but one polymorphic species, which shows vari-
ation in numbers of larval head-hairs and in mesonotal scaling, but no correlation
between these larval characters and the adults.” I cannot find such a statement by
either Shannon or Edwards. The latter author may have altered his views subse-
quent to his 1932 treatise. The assertion that there is no correlation between the
larval head hairs and the adult mesonotal scaling is incorrect, as shown in the present
revision, although the mesonotal scaling is subject to a certain amount of variation
in some species. Later Komp in unpublished notes, some of which at least were
made in the year 1951 and which were based mainly upon his collections in the
Panama Canal Zone in 1943-1945, identified specimens as thorntoni and podograph-
icus and also recognized 3 new species, alboapicus MS (this name used in the pres-
ent paper in the same sense), sp. A (=galindoi Schick) and sp. B (=zavortinki Schick).
Lane (1951:335; 1953:686-690), apparently after studying the type of homo-
eopus and perhaps | of the original males of argyrothorax at the USNM, took the
very dubious step of synonymizing these 2 species. This resulted in the scheme gen-
Schick: Terrens Group of Aedes (Finlaya) 7
erally accepted at the present: argyrothorax regarded as distinct, based upon its
unique male genitalia and having 1 synonym, homoeopus, and all the other nomi-
nal species lumped under terrens.
Knight and Marks (1952) followed the interpretation of species of Edwards (1932)
and did not make note of the changes proposed subsequent to that work. Forattini
(1965) followed the Lane classification (1953) but suggested that some of the geo-
graphic forms might represent full species. Finally, Belkin (1968:4) resurrected bra-
ziliensis from synonymy.
TAXONOMIC CHARACTERS
The characters used in this revision are discussed here. Unless otherwise indicated,
they do not show significant sexual dimorphism. They are arranged in the sequence
followed in the descriptions.
ADULTS
VERTEX. The decumbent scales of the vertex are usually silver or iridescent pur-
ple (referred to here as-“‘dark’’), rarely pale white, and usually narrow curved or
broad (flat), less frequently narrow spatulate. The broad scales are arranged in an
imbricate pattern and are appressed against the integument; the narrow curved scales
do not overlap, the underlying integument consequently visible between the scales,
and are not appressed against the integument. The narrow spatulate scales are us-
ually found in the median longitudinal line where they form a bilateral row of
anterolaterally directed scales overlapping along their lateral margins.
A median longitudinal line of silver and usually narrow curved scales which dif-
fer in color and/or form from the more lateral scales is present in most species. The
line may comprise a double or multiple row of scales. The scales laterad of the line
in most species are dark in the female and silver in the male, but there is no di-
morphism in the form of the scales; in a few species the more mesal scales are nar-
row curved and the more lateral scales broad.
OCCIPUT. The erect (forked) scales of the occiput may be pale, i.e. ivory colored
to pale brown, or dark, i.e. dark brown to black. Usually all or most of the scales
are pale in the female of most species but in some pale and dark scales may be pres-
ent in about equal numbers. All or most of the scales are dark in only a few species.
In gabriel the scalation varies from all pale through a mixture of pale and dark scales
to all dark and in buenaventura and aitkeni the median scales are pale and the lateral
scales are dark. The males of all the species except buenaventura and metoecopus
have all scales pale. The scales of buenaventura are like those of the female described
above and those of metoecopus are all dark.
PROBOSCIS. The length of the proboscis of the female is expressed relative to
that of femur I. The proboscis was measured from the apex of the clypeus to the tip
of the labellum and femur I from the basal articulation to the tip of the latero-
apical scales at a magnification of 45 X. The 2 structures were treated as being sub-
equal in length if they measured to within 0.04 mm of each other (i.e. to the near-
est quarter square of the net reticule described in the Introduction). The proboscis
showed 1 of 2 character states in most species, (1) shorter than or subequal in length
to the femur, or (2) longer than the femur.
PALPUS. The female palpus was not studied carefully. It appears, however, from
8 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
the few slides made, that the relative and absolute lengths of the penultimate seg-
ment afford specific characters.
In the male palpus, the length is expressed as the difference in the number of lab-
ellum lengths by which the palpus terminates from the tip of the proboscis. Measure-
ments were made primarily on those specimens in which the palpus was in the nor-
mal position, that is with its main axis diverging only slightly from that of the pro-
boscis (fig. 8). The measurements were accurate to 0.5 labellum length. There was
only a | labellum length variation in most species. The greatest variation, 3 labellum
lengths, occurred in argyrothorax, a species in which the palpus is unusually short.
The ventrolateral apex of segment 3 of the male palpus is provided with hairs
which vary in length and number. In some species only 1-3 hairs may be present
(typical thorntoni, fig. 9) and in others several, these forming a more or less well-
developed tuft (terrens, fig. 8). In those species with a tuft, the actual number of
hairs present may be of taxonomic value but this was not determined. The length of
the individual hairs or, when developed, of the tuft as a whole, may be divided into
2 taxonomic states, (1) shorter than or (2) at least as long as segments 4 and 5 com-
bined. Segment 4 is provided with a ventrolateral and a mesal row of hairs but only
the ventrolateral row is routinely treated in the descriptions since the development
of the 2 rows is correlated. The ventrolateral row is described simply as comprising
short or long, closely spaced or sparse hairs (see fig. 9, in which the hairs are short
and sparse, and fig. 29, in which the hairs are long and closely spaced). The develop-
ment of the hairs on segment 4 is more or less correlated with the development of
those of segment 3. :
MESONOTAL DISC. The mesonotal disc, which comprises the acrostichal and
dorsocentral areas, is provided with narrow curved scales that are either dark brown
- with a coppery luster (referred to here as “dark’’) or silver. When silver scales are
present they form more or less distinct markings that are not as intensely silvered as
that of the fossal macula (see below). Four types of markings are of taxonomic val-
ue, a transverse marking, an anterolateral strip, an acrostichal line, and a posterior
dorsocentral line. :
A transverse marking is present in many species but is usually developed only in
the male and in most of the species as an anterior band (fig. 17). The width of the
anterior band is expressed in terms of the level of the fossa (to the nearest quarter
or third) to which it is developed caudad along the dorsal midline. The band shows
intraspecific variation in width. The greatest variation occurred in male alboapic-
us, the band varying from 0.25 to 0.75 the length of the fossa, and in female thorn-
toni, being absent or present and varying from 0.33 to 0.75. The transverse silvering
of male insolitus, homoeopus and impostor usually extends into the posterior half
of the disc (figs. 29,31,33); the extent of this silvering along the longitudinal axis
may be subject to great variation (see homoeopus).
An anterolateral strip of scales bordering the fossa is developed only in those spec-
ies or individuals in which the fossal macula is coextensive with the fossa (see below)
(fig. 9). The strip superficially appears to be the mesal portion of the macula but its
scales lie on the dorsocentral area and are usually narrower than those of the macula
and are not imbricate. The extent to which the strip is developed, or even its pres-
ence or absence, is subject to variation in some species while in others it is sometimes
ill defined, being only represented by some scattered silver scales. The strip, as a
character, usually only complements that of the development of the fossal macula,
but shows a characteristic form in the male of sumidero (fig. 41).
An acrostichal line occurs in many species but may be present only in the male
Schick: Terrens Group of Aedes (Finlaya) 9
of some. The line may be (1) complete, extending from the anterior promontory
caudad to the prescutellar space (fig. 35), (2) incomplete, developed only at 1 end or
broken up into more than 1 segment by large areas of dark scales (fig. 29), or (3) ab-
sent. In some species the complete line is further characterized as being weak when
it is interrupted by a few dark scales.
A complete posterior dorsocentral line extends from the posterior tip of the fos-
sal macula caudad to the posterior margin of the mesonotum (fig. 41) and may be
weakly developed as in the case of the complete acrostichal line. The line is incom-
plete in many species, being developed only at the level of the prescutellar space
(fig. 11).
ACROSTICHAL SETAE. These are the setae of the acrostichal line that occur
caudad of those of the anterior promontory. The species generally fall into 3 classes
in respect to the acrostichal setae, (1) setae absent, (2) seta(e) present, the most
posterior occurring at about 0.25 to 0.5 from the anterior end of the acrostichal
line or (3) setae present, the most posterior occurring at about 0.5 or in the poster-
ior half of the line. In some of the species of the first class, 1 or more setae are oc-
casionally present but these usually occur within the anterior half of the line, rarely
as far caudad as 0.5. In some of the species of the second and third classes there is
some variation in regard to the placement of the most posterior seta, whether it be
in the anterior or posterior half of the line.
FOSSAL MACULA. The more or less well-defined anterolateral depression of the
mesonotum is treated as the fossa in this paper (fig. 25). The depression is usually at
least partially clothed with silver scales that are usually broad curved, imbricate and
appressed against the integument. These scales form the fossal macula. The macula
is coextensive with the fossa in many species, usually only in the male (fig. 17).
When coextensive, the mesal margin of the macula is convex since the mesal margin
of the fossa is also convex. In the remaining species the macula is variously reduced,
usually only on the mesal margin which may be convex (fig. 25), essentially straight
(fig. 52) or concave (fig. 37). The macula is greatly reduced in the females of a few
species and may be represented by a sublateral stripe (fig. 7) or a diffusely silver
scaled area or it may be complete suppressed (some insolitus).
The fossal depression is not distinct in some specimens. Whether or a the macu-
la is coextensive with the fossa in these cases can usually be determined by the cur-
vature of its mesal margin.
SUPRAALAR MACULA. The supraalar area is provided with a patch of appress-
ed, broad curved silver scales which form the supraalar macula. The macula, when
fully developed, is broadly joined to the fossal macula and caudally extends to the
most posterior seta. In the female of many species the macula is reduced anter-
iorly so that it is disjunct from the fossal macula (fig. 52). In some of the spec-
ies the macula may be reduced only anteromesally so that it is narrowly joined
to the fossal macula (fig. 31). The posterior margin of the macula is characteristically
truncate in a few species (fig.9). The supraalar macula of all the males of the Terrens
Group is broadly joined to the fossal macula.
PRESCUTELLAR SPACE. The females fall into 3 classes based upon the color of
the prescutellar scales, (1) at least some silver scales present, (2) silver scales present
or absent, (3) silver scales absent. The males usually have silver prescutellar scales,
but these may be lacking in soine species (galindoi, campana). The number of silver
scales tends to be greater in some species than in others but this was not carefully
Studied.
SCUTELLUM. The presence or absence of silver scales on the scutellum is gen-
10 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
erally correlated with the condition on the prescutellar space. The presence or ab-
sence of silver scales on the lateral lobe may show considerable intraspecific varia-
tion; consequently only the scalation of the midlobe is treated in the descriptions.
PLEURON. The presence or absence of a patch of scales on the subspiracular area
(ssp) is the only generally useful pleural scale character. This character is variable on-
ly in vargasi and in podographicus. The following 3 scale characters are of more lim-
ited taxonomic value: (1) scales of posterior pronotum (ppn) dark in aitkeni, pale in
other species; (2) scale patches of upper sternopleuron (stp) and mesepimeron (mep)
contiguous in buenaventura (fig. 25), not contiguous in other species; (3) mep patch
usually relatively small in species of the Thorntoni Subgroup, relatively large in diazi
and often divided into dorsal and ventral patches in terrens.
Many of the pleural hairs show considerable intraspecific variation in the inten-
sity of their pigmentation. Those of the prealar area (pra), however, were found to
be relatively constant in pigmentation and are useful for species discrimination.
These hairs are described as being pale or dark. When the hairs are dark, the inten-
sity of their pigmentation is compared to that of the hairs of the propleuron (ppl)
and sometimes of the upper mesepimeron (mep). Dark pra hairs are present in
many species, usually in both the sexes, but in some of the species the pra hairs
of the male may be pale. The pigmentation of the female pra hairs is variable appar-
ently only in galindoi.
LEG I. Femur I, and also femur II, are usually provided with a more or less well-
developed ventral silver line and a posterior patch of silver scales in the basal half
but only the posterior patch is of significant taxonomic value. The patch is described
here as being absent, weakly developed, small or well developed. A weakly developed
patch is one that comprises scattered silver scales; a small patch is more or less uni-
formly silvered but covers much less than one quarter of the area on the posterior
surface of the segment; a well-developed patch, the most common condition, is one
that is more or less uniformly silvered and covers at least one quarter of the area. The
patches on femur I and II may be subequal or dissimilar in size. In the latter case the
patch on femur II is usually larger than that of femur I.
A silver knee spot is present in only amabilis, the apical row of scales being sil-
vered.
The remaining silver markings, except probably for the exceptionally broad apical
silver band on tarsus 1-I in diazi (fig. 58), are too variable for taxonomic use.
LEG II. The silver ventral line of femur II is of some taxonomic value but since
its development is more or less correlated with that of the posterior silver patch it is
not treated in the descriptions. The posterior patch is discussed under Leg I.
The knee spot shows the following character states: absent, narrow, moderately
broad and broad, the latter being the usual state. These designations are explained
in the descriptions. The only marked intraspecific variation in the knee spot noted
in this study occurred in campana; the knee spot was absent in 3 of the 4 spec-
imens but was narrowly developed in the fourth.
The species generally fall into 3 classes in the development of a median dark
band on tarsus 1-II, (1) band incomplete, that is, not completely ringing the segment,
or complete and narrow, less than 0.33, (2) band complete and moderately broad,
about 0.33-0.5 or slightly greater, and (3) band complete and broad, about 0.6-0.75.
The markings of tarsus 2-II have limited taxonomic value. Most species show a
complete and broad apical dark band. In a few species the dark band is incomplete,
being longitudinally dissected by a streak of silver scales, or absent, the entire seg-
Schick: Terrens Group of Aedes (Finlaya) 11
ment being silver. The latter 2 states are usually developed only in those species in
which the median dark band of tarsus 1-II is incomplete or complete and narrow. In
diazi, however, the median dark band of tarsus 1-II is broad but tarsus 2-II is entirely
silvered.
Tarsi 3,4-II always lack silver scales and tarsus 5-II is silvered only in alboapicus
and occasionally in insolitus. |
LEG III. The widths of the bands on femur III and tarsus 1-IIIJ were measured
at a magnification of 45 X, and are expressed as the percentage of the total length
of the respective segments. Only the basal and subapical dark bands of femur III
and the basal and apical silver bands of tarsus 1-III of the females were treated.
The width of the bands in the males are similar to those of the females. The femur
was measured along its anterior surface from the basal articulation to the tip of the
lateral apical scales; the basal dark band from the basal articulation to the apical
notch of the band; and the subapical dark band from the basal notch of the dorsal
apex of the band (fig. 7). Tarsus 1-III was measured from the base of the scaled area
of the segment (represented usually by the base of the basal silver band) to the tip of
the apical silver scales; the basal silver band (actually an incomplete ring) at its
greatest width; and the apical silver band from its base to the tip of the apical scales.
The relative widths of the bands show characteristic ranges for the different spec-
ies but there is considerable overlap between many of them. The basal dark band of
femur III may be incomplete (the apical notch completely transecting the band) or
complete and as broad as 0.25 (nearly a quarter of the length of the segment); the
subapical dark band of femur III ranged from 0.11 to 0.49. The basal silver band of
tarsus 1-III may be absent or present and as broad as 0.20; the apical silver band
ranged from 0.07 to 0.38.
The width of the basal silver band of tarsus 2-III was not studied carefully but, ex-
cept for diazi in which it is unusually broad, the width appears to be correlated with
that of the apical silver band of tarsus 1-III.
Tarsi 3,4-III usually lack silver marking but in some species 3-III rarely shows a
narrow basal and apical band and 4-III a narrow basal band. Tarsus 5-III is silvered in
alboapicus and diazi but is dark in all the other species.
WING. Patches or lines of silver scales may be developed at the base of veins C,
R and Cu. The extent of these markings is subject to sexual dimorphism in most
species, the scalation being more extensively developed in the males. The scales on
vein C sometimes can be seen only from an anterior view.
Five character states may be recognized for the silver scalation of veins C and R:
(1) absent, (2) forming a small patch or short line, (3) a line reaching about 0.5 to
crossvein / (or to the level of h in the case of vein R), (4) a line reaching or nearly
reaching (or the level of h), and (5) a line (vein R only) extending well beyond
the level of h. The extent of the scalation of vein C shows some variation in the fe-
males of several species; some of the more extreme examples are found in metoe-
copus and zavortinki in which the states range from 1 to 3 and 2 to 4 respectively.
Variation in the males is less common, only 3 species showing occasional deviations
from the usual condition.
The silver scalation of vein R is of little taxonomic value in the females, states 1
or 2 being developed in most of the species. In the males, some species show only
state 1 while in others in which this is the usual state, state 2 may be developed.
Those typified by states 3 or 4 may show state 2 but not 1. The Maracay population
of the podographicus complex shows an extreme range of variation, from states 1
through 4.
12 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
Vein Cu is silvered only in the males of the Homoeopus Subgroup.
ABDOMEN. The silver scalation of the abdomen is apparently of little taxonomic
value. The width of some of the posterior silver sternal bands may be of some use
but this was not studied carefully.
MALE GENITALIA
SIDEPIECE. The length of the sidepiece is routinely given in the descriptions ©
principally to indicate the overall size of the genitalia. The sidepiece was measured
at a magnification of 100 X from the sternomesal base, at the point of the union
‘with the lateral lobe of the prosophallus, to the apex. There is considerable overlap
in the range of variation among many of the species but the modal value is some-
what useful for their characterization. The length varied from 0.25 to 0.46 mm.
A dense basal tergomesal tuft of long setae is present in several species (fig. 48)
and is a constant feature in these forms.
The median sternomesal area shows 3 specialized structures of taxonomic value, a
convexity, a sclerite and a tuft. The median sternomesal convexity is a broadened re-
gion of the sternal flap and may be absent or weakly developed (fig. 52), moderately
developed (fig. 44) or strongly developed (fig. 31). This structure is of limited taxo-
nomic value serving chiefly to characterize homoeopus in which it is usually stOte
developed.
The median sternomesal sclerite is a rounded and dorsal extension of the stone
flap and when present is developed at the apex of the median sternomesal convexity.
The sclerite may be absent, weakly developed (fig. 52) or well developed (fig. 31).
The median sternomesal tuft originates on the median sternomesal sclerite and
comprises fine setae (fig. 31). The tuft is present in all the species with a well-devel-
oped median sternomesal sclerite and in some of the species with a weakly developed
sclerite. The tuft of galindoi and campana differs from the other species in that the
setae are more numerous, more closely spaced and wavy (figs. 44,46). |
In addition to the above features of the sidepiece, a rather inconspicuous but dis-
tinctive subapical tergomesal tuft of very fine setae is present in argyrothorax (fig.
13).
PROSOPHALLUS. Belkin (1968:9) proposed the term prosophallus for the stern-
al structure between the base of the sidepieces of the male genitalia of the Dixinae.
He suggested that the prosophallus was homologous with the claspette of the Culici-
nae. The prosophallus is interpreted here as comprising the interbasal fold and clasp-
ette in the sense of Freeborn (1924:205) and subsequent authors.
The measurements of the prosophallus were made at a magnification of 210 X as
shown in figure 9. The length, primarily a measure of the length of the stem, varied
from 0.07 to 0.15 mm. The intraspecific ranges of variation overlap considerably
but the length is still of use in the characterization of some species. The width is
used in this paper only to distinguish zavortinki from other species of the Terrens
Subgroup.
The interbasal fold shows 3 basal lobes and a pair of distal lobes, the latter termed
mesal lobes (fig. 9). The basal lobes are of taxonomic value for the characterization
of galindoi. In this species the median lobe projects farther cephalad than the lateral
lobes (fig. 44). The median lobe in the other species projects cephalad to about the
same level as the lateral lobes (fig. 46). The mesal lobe varies among the species in
the inclination of its distal margin. The lateral portion of the mesal lobe may be de-
clivous mesad (fig. 31), not appreciably inclined from the horizontal (fig. 17) or de-
Schick: Terrens Group of Aedes (Finlaya) 13
Clivous laterad (fig. 13). The latter condition occurs only in argyrothorax. The angle
of inclination from the horizontal was measured in those species in which the lobe is
declivous mesad. Measurement was made with the aid of an ocular net reticule (see
Introduction) at a magnification of 860 X and is given in units of 15° in the des-
criptions. The angle ranged from 0 to 45°; the usual intraspecific variation was less
than 15° but in many species there are occasional specimens which increase the var-
iation to 15° and rarely 30°.
The stem is of particular importance in the characterization of the Podogtaphicus
Subgroup in which it is characteristically bowed (fig. 52). This type of stem is oc-
casionally developed unilaterally and rarely bilaterally in the other species. The
stems, in those species in which the central axis is not appreciably curved, may be
divergent (fig. 27), parallel (fig. 17) or convergent (fig. 13). Two or 3 of these states
may be shown by | species but usually only 1 is typical of that species.
The filament in all the species, except argyrothorax (fig. 13), is in the form of a
stout seta whose apical portion is curved to form a hook (fig: 8). The length of
the filament was determined at a magnification of 210 X on whole mounts of the
genitalia. It was divided by the length of the prosophallus to determine the fila-
ment ratio which is given to the nearest 0.05 in the descriptions. The ratios ranged
from 0.30 to 1.35 and the usual intraspecific variation was 0.25. The hook may
be strongly angulate (fig. 31), weakly angulate (fig. 27) or evenly curved (fig. 29),
the latter 2 conditions referred to simply as not strongly angulate in the descrip-
tions. In most species the hook is either strongly angulate or not strongly angu-
late. A strongly angulate hook is occasionally developed in some of the species
in which the hook is usually not strongly angulate; it seems that at least in some
of these cases the strong angularity is the result of distortion produced in the prep-
aration of the slide. Aedes homoeopus, on the other hand, seems to show actual
intraspecific variation, the usually strongly angulate hook being weakly angulate
in a few specimens. :
AEDEAGUS. The aedeagus is of little taxonomic value. It is relatively narrow in
buenaventura (fig. 25) and the lateral sclerotized portion in argyrothorax is narrow-
er than in the other species. The absolute length is of limited use. Measured at a
magnification of 210 X, the length varied from 0.10 to 0.16 mm. The intraspecific
variation was usually 0.02, rarely as great as 0.03 mm.
OTHER STRUCTURES. The other parts of the male genitalia are apparently of
limited taxonomic value. In argyrothorax (see) tergite IX, the clasper and paraproct
Show unique features and in gabriel the spiniform of the clasper appears to be rela-
tively longer than in the other species.
PUPAE
ORNAMENTATION. The mesonotum and sometimes the legs cases and the mid-
dle portion of the wing case are more heavily pigmented than the remainder of the
cephalothorax. A pale inverted V-shaped marking is formed in those specimens in
which all 3 of these structures are heavily pigmented. This marking extends from
the base of the mouthparts case caudolaterad through the base of the wing case
(indicated in part in fig. 9) and may be weakly differentiated in those specimens
in which the wing case pigmentation is weakly developed or does not extend across
the entire width of the case.
CHAETOTAXY. Only 2 hairs were found to be of general taxonomic value. Hair
I-I varied in the predominant number of secondary branches per primary branch
14 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
and ranged from predominantly single to multiple. The range of variation is restrict-
ed in most species but in others the primary branches range from predominantly
single to predominantly multiple.
Hair 2-II shows taxonomic value in its position relative to 3-II along the trans-
verse line. The position can vary significantly among the species but in regard to
the usual position most species show 1 of the following character states, (1) 2-Il
always laterad of 3-II, (2) 2-II laterad of 3-II or mesad of 3-II for less than 0.3 the
distance from 1-II to 3-II, or (3) 2-II mesad of 3-II for 0.3 or more. A fourth char-
acter state, mesad for more than 0.3 may apply to 1 or 2 species but only a few
specimens of these species were available. The distance of 2-II mesad of 3-II was
determined to the nearest tenth with the aid of the ocular reticule at powers rang-
ing from 100 X to 210 X.
PADDLE. Three paddle characters are of general taxonomic value, the shape of
the apex, the general pigmentation and the pigmentation of the ventral midrib. The
apex may be weakly emarginate (fig. 39), evenly rounded (fig. 9), or weakly to
strongly produced (fig. 40). In many of the species the apex is not at all produced
or at most only weakly produced. In other species the apex is usually produced but
the degree of the production is variable. The extremes are illustrated in the pupal
drawings of some of the species.
In several species the paddle is clear, lacking all apparent pigmentation. In the
other species the paddle is pigmented to various intensities and the amount of pig-
mentation shows considerable intraspecific variation; the paddle of some specimens
is very weakly pigmented but this condition can usually be differentiated from that
of the clear paddle by the slightly darker overlapping portions.
The ventral midrib is pigmented only in species with a generally pigmented pad-
dle. In most of these species the midrib is weakly pigmented along most of its
length and does not contrast strongly with the rest of the paddle; the other species
show a moderately to strongly pigmented midrib which contrasts rather strongly
with the rest of the paddle. The apical portion of the midrib is more weakly pig-
mented than the remainder in most species but in some of the species with a mod-
erately or strongly pigmented midrib the pigmentation may be as intense (fig. 31)
or even more intense (fig. 15) in the apical portion.
LARVAE
CHAETOTAXY. The following hair characters were found to be of greatest tax-
Onomic value: the relative lengths of 7,11,14-C, 11-P, 8-T, 5-I and 5-VII; the num-
ber of branches of 5,6,14-C, bmh, 1-A, 14-P, 3,4-M, 3,10-III, 3-VI, 4,10,12-VII
and 2-VIII; and the position of 2-II and 5-VII.
The length of hair 8-T is given in terms of the length of the metathoracic pleural
tubercle. Measurements were made at a magnification of 210 X and only in those
specimens in which the tubercle was oriented in a reasonably horizontal manner.
The tubercle was measured from its base to the apex of the longest distal spine. The
lengths of the other hairs, also measured at a magnification of 210 X, are expressed
relative to either the mental plate or to a particular hair.
The number of branches of hairs 7-C, 1,4,5,7-P are routinely given in the species
descriptions as supportive data and to characterize different populations of particu-
lar species.
COMB SCALES. The species may be roughly divided into 3 classes in respect to
the range of variation in the number of comb scales, which is rounded to the near-
Schick: Terrens Group of Aedes (Finlaya) 15
est 10: (1) 20 to 40, (2) 30 to 50, (3) 50 to more than 100. Six species fall into the
first 2 classes and the remaining into the third class.
The most useful comb scale character is the absolute length of the free portion
of the midapical scale. This portion was measured from the most distal part of the
sessile portion, appearing as a crescent in a slide mount, to the apex (excluding the
fringe) at a magnification of 430 X. The midapical scale occasionally assumes an
anomolous broad form that differs markedly from the adjacent scales; this type was
not measured. The free portion ranged from 0.017 to 0.052 mm and usually showed
an intraspecific variation of about 0.009 mm;; in insolitus the variation was unusually
great, 0.025 mm, due in part to geographic variation.
The length of the free portion of the most apical scale relative to that of the ses-
sile portion provides a character supplementary to that of the absolute length of
the free portion. The length is expressed simply as being shorter than, subequal in
length to or longer than the sessile portion.
Two other characters of limited use are not treated routinely in the descriptions.
One is the number of rows of comb scales, varying from 3 to 9 among the species.
Since the usual range of intraspecific variation is 3 rows, this character has little
general value. It is useful, however, in separating galindoi from campana and tehuan-
tepec from schroederi. The second character is the shape of the free portion of the
scales. In most species, except sometimes for the basal scales, the free portion is
spatulate or pandurate. In some daryi, however, the comb comprises only ligulate
or a mixture of ligulate and almost awllike scales (fig. 51).
SIPHON. Three siphon indices are routinely cited in the descriptions but have
limited general diagnostic value since the intraspecific ranges of variation overlap
to a considerable extent. The modal values, however, provide supportive characters.
All measurements were made at a magnification of 100 X.
Relative siphon length (L/S). This is the length of the siphon (L) divided by the
length of the saddle (S). The siphon was measured along its main axis from the dor-
sal base to the dorsal apex (fig. 10); the saddle was measured along its middorsal line.
The L/S ratio'was proposed by Colless (1957:87) as a means for deriving a satis-
factory relative siphon length in specimens of all conditions, from “‘badly crushed
skins to whole larvae,” and is used here since so much of the material on hand con-
sists of skins with the siphon crushed to various degrees. Colless (1962:364) pre-
sented data on Culex (Lophoceraomyia) comparing the L/S and classical siphon in-
dex; the L/S showed less intraspecific variability and a better discriminant function
in interspecific analyses. The L/S in the Terrens Group showed a range of 1.82 to
2.86 and a usual intraspecific variation of 0.33 to 0.38.
Pecten row length index (P/L). This is the distance of the most apical pecten
tooth from the dorsal base of the siphon (P), measured along the same line of pro-
jection as for the siphon length, divided by the siphon length (L) (fig. 10). The pec-
ten was measured on both sides of the siphon and the average of the 2 was used
for the index. The P/L showed a range of 0.44 to 0.64 and a usual intraspecific var-
iation of 0.08.
Hair 1-S placement index (H/L). This is the distance of the alveolus of hair 1-S
from the dorsal base of the siphon (H), measured along the same line of projection
as for the siphon length, divided by the siphon length (L) (fig. 10). The position of
the alveolus was measured on both sides of the siphon and the average of the 2 was
used for the index. The H/L showed a range of 0.51 to 0.69 and a usual intraspecific
variation of 0.06 to 0.08.
SADDLE. The saddle offers few characters of diagnostic value. In most of the
16 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
species the saddle extends at least halfway, or a somewhat greater distance, around
the anal segment and the ventral margin usually shows a rounded notch of vari-
able breadth or is irregularly undulate. Three species show deviations from this
type of saddle. In argyrothorax the saddle extends less than halfway around the
segment (fig. 14). In tehuantepec and schroederi (Tehuantepec Subgroup) the sad-
dle extends much farther around the segment than in the other species (figs. 59,
60), although the exact extent is somewhat variable, and a narrow marginal or sub-
marginal slit is developed at the ventral margin of the saddle. The slit is submarginal
in those specimens in which the saddle is developed to a particularly great extent
ventrad and marginal in those in which the saddle extends ventrad to a lesser extent.
A few of the specimens with a submarginal slit show a variously developed strongly
sclerotized band around the slit (fig. 60).
GILLS. The length of the gills, relative to that of the anal saddle measured along
its dorsal midline, shows considerable intraspecific variation and is of little taxonom-
ic value. The gills in a few species are short, less than the length of the saddle but
even in these the gills may be much longer than the saddle. The length of the gills
tends to be relatively stable, however, in a particular collection.
SYSTEMATICS AND EVOLUTION
The Terrens Group is most easily characterized by the leg banding of the adults.
Typically, tarsi 1-II,III show a basal and apical silver band and tarsi 2-II,III only a
basal silver band while the remaining segments are entirely dark; in 2 species, how-
ever, the fifth tarsal segment of at least leg III is silvered. Aedes buenaventura is
unique in the group in the absence of the midtarsal and hindtarsal silver bands ex-
cept for a very narrow one at the base of 1-II. The adults of this species can be read-
ily separated from other New World Finlaya which lack prominent silver tarsal bands
by the broad dark basal band of femur III, a character developed only in the Terrens
Group; also the mesonotal disc of the male shows an anterior silver band and the
hairs of the pra are dark, features of the Terrens Group that are not usually shown
by other New World Finlaya.
The pupae are generally similar to other Finlaya of the region, but the trumpet is
more darkly pigmented than in most of these other Finlaya.
The larvae of the Terrens Group can be distinguished from the other Finlaya, ex-
cept kKompi and scutellalbum, by an at least moderately large patch of usually spat-
ulate or pandurate comb scales with an even fringe of fine spicules. Aedes kompi
can be separated from the species of the Terrens Group in the larval stage by the
usually stronger spine of the metathoracic pleural tubercle and by the usually ab-
ruptly tapered distal portion of the siphon, this portion of the siphon being more
or less evenly tapered in the Terrens Group; and scutellalbum by the more spiculose
anal saddle and the stouter hair 2-A. »
The Terrens Group of the present revision corresponds to Group B (TERRENS-
group: GUALTERIA), Subgroups I and II (terrens and argyrothorax) of Knight and
Marks (1952:522-523). Subgroup I of these authors included terrens and most of the
other nominal species of the group and Subgroup II only 1 species, argyrothorax.
Four characters separated Subgroup II from I: (1) the broad scales of the vertex (pre-
sumably referring to those along the longitudinal midline), (2) the short male pal-
pus, (3) the short, stiff hairs of segments 4 and 5 of the male palpus and (4) the
broadly expanded claspette filament. Aedes thorntoni, not included in the scheme
Schick: Terrens Group of Aedes (Finlaya) 17
since the authors had not seen the male, apparently is an annectent form. This spec-
ies shows character states 1 and 3 of Subgroup II and a relatively long male palpus
and simple claspette filament as in Subgroup I. Aedes thorntoni and argyrothorax
actually show many characters that indicate that they are closely related and it
seems best to treat them together as a single group (see Thorntoni Subgroup). Since
state 3 of Subgroup II is shown by some of the new species of the present revision
that would fall into Subgroup I, only the first character state of Knight and Marks
would remain separating the 2 subgroups. Subgroups I and II do not appear to rep-
resent primary subdivisions of the Terrens Group and consequently are not recog-
nized in the present revision. The subgroupings within the group is discussed below.
Three of the new species of the present paper do not key to Group B of Knight and
Marks. Based upon leg markings, alboapicus and diazi key to Group C (LONGIPAL-
PIS-group) and buenaventura to Group H (GENICULATUS-group). The pattern of
leg markings would seem to be a dubious group character.
Twenty-eight species are formally recognized in the present revision. Some of
these are rather broadly interpreted and may actually represent species complexes.
In addition, there are at least 7 more species which are not formally treated here
either because of their representation by inadequate material or because they came
to my attention when the manuscript was in its final stage of preparation. Two of
these species (and probably more, as indicated under ferrens) are from northeastern
South America, 2 from the Andes in Ecuador and 3 from northwestern Argentina.
The previous interpretation, that terrens itself was a highly variable and widespread
species, is completely invalidated by an analysis of the morphology of the different
stages and the distribution of the forms; also, there is correlation between adult,
male genitalic, pupal and larval characters. Sympatry of species is common and in
several cases 2 species have been collected in the same treehole.
The species in this revision are based primarily on adult scale patterns and other
adult non-genitalic features but often the male genitalia and usually the larvae pro-
vide important supportive taxonomic data. The male genitalia of many of the spec-
ies show only subtle differences which are of a statistical nature and consequently
a key based on genitalia is not included here. The pupae offer the fewest characters
of taxonomic value and in general are too similar and variable to be placed in a key;
some of the species, however, have a distinctive chaetotaxy and the paddles often
can be used for separating species or subgroups.
Fourteen primary subgroups are recognized in the Terrens Group in the present
revision. They are characterized below in those stages that show unique or unusual
features. For more complete characterizations see the individual subgroups in the
taxonomic treatment.
Thorntoni Subgroup. Vertex of adults with decumbent scales along longitudinal
midline broad. Larval hairs 7,11-C, 8-T and 5-I unusually short.
Bertrami Subgroup. Known only in the female. Mesonotal disc with broad anter-
ior silver band. :
Terrens Subgroup. A varied group showing no apparent unique features.
Alboapicus Subgroup. Tarsal segments 5-II,III of adults silvered. Pupal hairs 5-C,
1-P unusually long.
Buenaventura Subgroup. Adults with scale patches of upper stp and mep contig-
uous; mid- and hindtarsi with silver band only at base of 1-II. Sidepiece of male gen-
italia with specialized sternal subapical seta. Larva with hair 5-VII cephalad of 4-VII.
Metoecopus Subgroup. Erect scales of vertex of male dark.
Insolitus Subgroup. Mesonotal disc of males usually largely silvered.
18 Conti ta mieesBikt Inatiivols Sato. 31970
Aitkeni Subgroup. Known only in the female. Fossal macula with pale golden
scales and ppn with dark scales.
Homoeopus Subgroup. Mesonotal disc of males usually largely silvered; veins R
and Cu of males with unusually long basal line of silver scales.
Heteropus Subgroup. A varied group showing no apparent unique features.
Galindoi Subgroup. Larval hair 2-II situated relatively far laterad.
Podographicus Subgroup. Claspette stem of male genitalia characteristically bow-
edc:
Tehuantepec Subgroup. Anal saddle of larvae relatively more complete than in
other groups and with marginal or submarginal ventral slit.
Diazi Subgroup. Known only in the female. Mid- and hindtarsi shaggy and with
silver bands of forelegs unusually broad.
I have not been able to assemble these primary units into larger groups character-
ized by correlated features in the adults and immature stages. Possibly the first 2 to
4 subgroups listed could be combined but this would result in a disproportionately
large and varied group. Some of the difficulties involved in recognizing larger groups
will become apparent in the discussion that follows on the evolution within the
Terrens Group.
There are possibly 2 major phyletic lines in the Terrens Group and for the pur-
pose of discussion these are termed the Terrens and Podographicus lines. The Ter-
rens line comprises the following subgroups, Thorntoni, Bertrami, Terrens, Alboapi-
cus, Buenaventura, Insolitus and possibly Metoecopus; and the Podographicus line,
Homoeopus, Heteropus, Galindoi, Podographicus, Tehuantepec and possibly Diazi.
The Aitkeni Subgroup cannot be satisfactorily referred to either line.
Fifteen characters are of use in distinguishing these lines. They are given below,
the Terrens character first and the Podographicus character second; an asterisk after
a character indicates that it is shown by all the members of the line. The Metoe-
copus Subgroup is not included in the characterization of the Terrens line since it
shows very few of the Terrens characters.
Adults. (1) Mesonotal disc of males: anterior silver band present—anterior silver
band absent; (2) fossal and supraalar maculae of females: broadly joined—not or nar-
rowly joined; (3) pra hairs of females: dark—pale; (4) basal dark band of femur III:
broad—incomplete or narrow; (5) silver scalation at base of vein C of males: line
reaching crossvein h*—small patch or line reaching at most 0.5 to h®*.
Pupae. Cephalothorax: pale inverted V-shaped marking present—V-shaped mark-
ing absent*.
Larvae. (1) Hair 3-M: branched—single; (2) 4-M: triple or multiple—single or doub-
le*; (3) 3-III: triple or multiple—single or double*; (4) 10-III: branched—single*;
(5S) 3-VI: branched—single*; (6) 4-VII: branched*—single*; (7) 10-VII: branched—
single; (8) 12-VII: branched—single*; (9) 2-VIII: branched —single.
The subgroups and species of the Terrens line vary in the number of the Terrens
line characters that they possess. In the Thorntoni Subgroup all 15 of the Terrens
characters are shown by thorntoni and 14 of the 15 by argyrothorax; most are
shown by the Terrens, Alboapicus and Insolitus Subgroups but only 6 by the Buena-
ventura and 3 by the Metoecopus Subgroups. On the other hand, all the species of
the Podographicus line show either all or a majority of the Podographicus line char-
acters. Each of the 15 character states for the Terrens line represents either a posi-
tive character state, that is, a more extensively silvered or more heavily pigmented
condition, or a state of greater number of elements (hair branches) and those of
the Podographicus line either a negative character state or a state of a lesser number
Schick: Terrens Group of Aedes (Finlaya) 19
of elements.
It is suggested here that the Podographicus condition is the derived one. The pres-
ence of Podographicus character states in different combinations among species of
the Terrens line seems to be most logically explained by a process of independent
reductions of the Terrens states to those of Podographicus. One Podographicus char-
acter in particular, the narrowly joined or disjunct fossal and supraalar maculae of
the females, is strongly suggestive of a derived state since the maculae are broadly
joined in all the males of the Terrens Group. Two other characters suggestive of a
derived state are the narrow or incomplete basal dark band of femur III and the
short line or small patch of basal silver scales on vein C of the males; these markings
are more extensively developed in the Terrens line and are variously and more poor-
ly developed in most of the species of the Podographicus line.
Conversely, it seems less likely that the Podographicus condition is the more
primitive since the presence of Podographicus character states among the species
of the Terrens line would have to be explained either by (1) the acquisition of the
positive Terrens states from the negative Podographicus states independently among
the species or (2) the derivation of the Terrens condition from the Podographicus
condition followed by independent reversions to the Podographicus states. Also, as
indicated above, at least some of the Podographicus states appear to be derived.
Since, in the evolution of the Terrens line, there were apparently transformations
of Terrens to Podographicus states independently among the species, it is possible
that more than 1 phyletic line showing the Podographicus condition exists which
cannot be detected because of the parallel evolution. One possibility is the existence
of 2 major lines, 1 represented by the subgroups that show acrostichal setae (Homo-
eopus, Heteropus and Galindoi) and those in which these setae are absent (Podo-
graphicus, Tehuantepec and Diazi). The position of pupal hair 1-II in relation to 3-II
supports this scheme to some degree but there are apparently no other characters
to substantiate it.
The distribution of the subgroups provides the following data in relation to the
phyletic lines. In the Terrens line, the Thorntoni, Bertrami, Terrens and Alboapicus
Subgroups (the “‘core’”’ of the Terrens line) occur primarily in the Atlantic lowlands
of Central America and in the Atlantic drainages of South America; some of the
species also cross over to the Pacific lowlands of eastern Panama at the Canal Zone;
the Insolitus Subgroup occurs in the highlands of Central America and northern
South America and the Buenaventura and Meteocopus Subgroups, which are the least
Terrens-like of the line, are found in the Pacific lowlands of northern South Amer-
ica. The subgroups of the Podographicus line occur in the highlands and/or low-
lands of Mexico and Central America; 1 form belonging to the Podographicus Sub-
group, occurs disjunctly at moderately high elevations in northern Venezuela. In
relation to the possible diphyletic composition of the Podographicus line discussed
above, the Podographicus and Tehuantepec Subgroups (but not the Diazi Subgroup)
are distributed primarily in the coastal lowlands of Central America while the re-
maining subgroups are found primarily at higher elevations.
In summary, the Terrens line appears to be the more primitive of 2 hypothet-
ical major phyletic lines. This line is primarily Panamanian and South American in
distribution. The Thorntoni, Terrens and Alboapicus Subgroups, and possibly Ber-
trami, form the ‘‘core”’ of the Terrens line and show a similar type of distribution.
The Insolitus Subgroup is morphologically similar to these subgroups but shows
a different type of distribution. The Buenaventura Subgroup and especially the
Metoecopus Subgroup show a tenuous relationship with the preceeding subgroups
20 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
and occur in a different geographic area. The remaining subgroups, except possibly
for Aitkeni, comprise the Podographicus line which may actually be polyphyletic.
This line is primarily Mexican and Central American in distribution.
The Insolitus Subgroup is possibly the most primitive subgroup of the Terrens
Group. This is suggested by the following rather meager data and assumptions. (1)
The Insolitus Subgroup is represented in the area which seems to be the most likely
primary center of distribution of the Terrens Group, southeastern Central America
(and possibly northeastern South America), by disjunct and distinctive montane pop-
ulations. (2) The subgroup belongs to the apparently primitive Terrens phyletic line
and also shows the following 2 characters that perhaps link it with the Podographi-
cus line, the largely silvered mesonotal disc which is developed elsewhere only in the
Homoeopus Subgroup and the relatively long hair 11-P of the larva which is long
elsewhere only in heteropus. and the Galindoi Subgroup. (3) If the largely silvered
mesonotal disc of the Insolitus males is the primitive condition, then the anterior
silver band of the disc of most of the remaining subgroups of the Terrens line could
be the product of a caudal reduction (see homoeopus). I would suggest that a
southern population of an ancestral Insolitus stock gave rise to the Terrens Sub-
group. The latter may be the most primitive of the southern subgroups since some
of the species (zavortinki and apollo) appear to be relatively nonspecialized and
since, as treated here, it comprises a diverse array of species. The Thorntoni and
Alboapicus Subgroups appear to have been derived from the Terrens Subgroup;
they share with the latter many characters of subgroup significance but also show
some apparently specialized features. The Buenaventura Subgroup and the appar-
ently unrelated Metoecopus Subgroup probably represent lines derived indepen-
dently of the Terrens Subgroup since they occur in the Pacific rather than the At-
lantic drainage of South America. I would also suggest that the northern highland
Aitkeni Subgroup and subgroups of the Podographicus line which show acrostichal
setae arose from 1 or more northern populations of the ancestral Insolitus stock;
1 species of these subgroups, daryi, is now represented by widely disjunct popula-
tions in Guatemala and southeastern Panama which suggests a differentiation in an
area located between these ends of Central America. The Podographicus Subgroup,
which lacks acrostichal setae, possibly differentiated in northeastern South America,
perhaps from a species of Insolitus or from some member of the Terrens Subgroup; if
the subgroup did evolve in this area it must have spread northward and crossed
into the Pacific drainage. The Isthmus of Tehuantepec perhaps was the passageway
to the Pacific since distinct populations of podographicus now occur to the north
and to the south of the Isthmus on the Pacific side of Middle America. If on the
other hand, the Podographicus Subgroup differentiated somewhere in its present
lowland Central American range, its anomolous presence at moderately high eleva-
tions in Venezuela must be explained.
Although this phylogenetic scheme is only tentative it at least provides a work-
ing hypothesis. A less speculative phylogenetic scheme cannot be formulated until
the fauna of the Atlantic lowlands of Central America and the montane faunas of
southeastern Central America, northeastern South America and the Andes are bet-
ter known.
DISTRIBUTION
The Terrens Group is distributed from the northernmost limits of the Neotropical
Schick: Terrens Group of Aedes (Finlaya) 21
region (in the sense of Maldonado-Koerdell 1964:20, i.e. from the state of Nayarit
in the West, across Mexico, including the Mesa Central, to the southern limits of
the state of Tamaulipas in the East) southward into Ecuador, Peru, Bolivia, north-
ern Argentina and southeastern Brazil and at elevations from sea level to at least
8,000 feet (fig. 1).
It is of interest that the Terrens Group is absent in the West Indies. Maldonado-
Koerdell, in his treatment of the geohistory of Middle America (1964:15-18), indi-
cated that northwestern Central America (southeastern Mexico to northern Nicar-
agua) and the West Indies (at least in part) formed a single land mass, Caribbean ~
Land, from the early Tertiary through a major part of the Miocene. Southeastern
Central America was broken up into several (or many) islands at this time. It is my
premise that the Terrens Group originated on these islands; this is based upon the
following tenuous data: (1) the greatest diversity of species occurs in this area,
(2) several relict montane species or populations of widespread species occur here
and (3) the area represents the center of the range of the group. Uplift of south-
eastern Central America occurred in Miocene-Pliocene times and the Terrens Group
spread northward and southward. The absence of the group in the West Indies may
be explained by the dispersal of at least the low elevations species into northwestern
Central America after the subsidence of the transoceanic portion of Caribbean Land
in the late Miocene.
The distributions of the subgroups are illustrated in figures 2-6. Half of the 14
subgroups are monotypic and show restricted geographic ranges. Four of the poly-
typic subgroups, Homoeopus, Heteropus, Galindoi and Podographicus, show striking
discontinuities in their ranges. Two of the subgroups, Thorntoni and Terrens, are dis-
tributed primarily in the Atlantic lowlands of Central America and in the Atlantic
drainages of South America, but some of the species cross over through the Canal
Zone to the Pacific side of eastern Panama; these species have not been recorded west
of the Canal Zone on the Pacific side. Of interest, podographicus, the common spec-
ies of the Pacific lowlands to the northwest and north, apparently does not occur in
southeastern Panama although it is recorded from the province of Panama west of
the Canal Zone (see figs. 2,3,6).
BIONOMICS
The species of the Terrens Group are principally sylvan but some of the species
occur in populated areas. Bonne and Bonne-Wepster (1925 :424) took 1 of the origi-
nal males of argyrothorax in a house in Paramaribo; Prosen, Carcavallo and Martinez
(1964:103-104) reported ‘“‘terrens” adults in rural areas of Bolivia; and homoeopus
larvae were taken on the grounds of the University of Costa Rica and in Parque
Bolivar in San Jose, Costa Rica (UCLA collections). On the other hand, Shannon
(1931a:23) observed that terrens did not occur in the more or less natural areas of
suburban Salvador, Brazil; terrens larvae were not taken in numerous treehole col-
lections although Culex conservator larvae were abundantly represented.
The vertical distribution of terrens (sens. lat.) has been investigated by Galindo,
Trapido and Carpenter (1950), Galindo, Carpenter and Trapido (1951) and Trapido,
Galindo and Carpenter (1955a,b) in their studies on diurnal forest mosquitoes, car-
ried out in different parts of Panama. The terrens of these authors almost certainly
included campana and probably thorntoni, zavortinki and podographicus, and pos-
sibly aitkeni and other species. In the 1950 and 1955a studies, biting-landing collec-
Za Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
tions were made on the ground and in the canopy; most specimens were taken on
the ground, about 68% of the 25 specimens in the 1950 study and about 83% of
the 1533 specimens in the 1955 study. Bamboo larval traps were used in the 1951
and 1955b studies and were placed near ground level, at 20-30 feet (low platform)
and at 40-50 feet (upper platform); in the 1951 study it was noted simply that
most larval collections occurred in the traps situated near the ground but that on
several occasions larvae were found in the traps at the lower and upper platforms;
and in the 1955b study 80% of the 252 larvae were taken near the ground, and the
remainder in the canopy. The ground and canopy mosquitoes possibly represent
different species.
All of the larval collections of the present project were made at heights from be-
tween ground level to 15 feet, presumably too small a range to indicate height
specificity.
Flight range figures for terrens in the state of Minas Gerais, Brazil, were obtained
by Causey, Kumm and Laemmert (1950). Adults were recaptured 4.3-4.4 km from
a point of release after 1-2 days and 2.2-5.6 km after 1-17 days in 2 separate experi-
ments.
Many of the species have been observed to be anthropophilic. Those definitely
known to bite man are insolitus in Trinidad, homoeopus from Cordoba, Mexico and
idanus (UCLA collection data) and what is probably campana (Galindo, personal
communication). The following species have been taken in biting-landing collections
but the data do not indicate whether they were actually biting: amabilis, apollo,
berlini, bertrami, metoecopus, sumidero, terrens from Rochina, Brazil, vargasi (UCLA
and other original collection data), argyrothorax in Brazil (Causey, Causey, et al,
1961:243), and what may be braziliensis (Kumm and Novis, 1938:502).
Davis and Shannon (1931:716) and Shannon, Whitman and Franca (1938:110)
fed terrens from Brazil on monkeys and Kumm and Novis (1938:502) fed what may
be braziliensis on Cebus monkeys and on agoutis.
All data on the activity of females indicate that it is diurnal. Fisher and Verity,
however, observed suwmidero flying in a shelter at night (MEX 115-2).
Egg deposition by ‘‘terrens’’ was observed by Galindo, Trapido and Carpenter
(1951:121). The eggs were laid about 1 inch above the water level in a treehole.
Galindo, Carpenter and Trapido (1955:162), using bamboo containers containing
organic debris, showed that the hatching of the greatest number of eggs occurred
after the first flooding. A substantial number hatched after the fourth and fifth
flooding cycle and the last at a ninth cycle.
The larvae breed primarily in treeholes. Many species have also been found in cut
or broken bamboo and in artificial containers and it seems likely that all or most
species can utilize these habitats. Also, 1 collection of gabriel was made in a volcanic
rockhole and 1 of insolitus in a coconut shell. The size of the treehole in general
does not appear to be a critical factor in breeding site preference, most species be-
ing recorded from both “small” and ‘“‘large’ treeholes in our collections. How-
ever, alboapicus and metoecopus might show such a preference since only small
treeholes or bamboo have yielded these species. Galindo, Carpenter and Trapido
(1951:110,111) obtained “‘terrens” principally in their bamboo traps with large
Openings (apparently their open-top trap) and very few in their traps with a lid
and a one-half inch opening.
Schick: Terrens Group of Aedes (Finlaya) 23
DISEASE RELATIONS
Although probably most of the species are anthropophilic none has been implicat-
ed in disease transmission. Most experimental work has been carried out on terrens in
Brazil. Davis and Shannon (1931:716) observed that Yellow Fever virus remained
active in terrens from Salvador, Brazil, for at least 2 weeks and that a monkey was
killed by the injection of a single specimen. However, since the mosquitoes were
reluctant to feed for a second time in the laboratory and since they were not cap-
tured in the forest on either human or animal bait, the authors concluded that it
was improbable that terrens was a factor in Yellow Fever transmission. Shannon,
Whitman and Franca (1938:111) fed terrens, collected in the forest of the state of
Rio de Janeiro where Yellow Fever had been reported, on infected monkeys. Only
58 of the 503 mosquitoes fed and none of these was positive for the virus.
TAXONOMIC TREATMENT
Terrens Group
ADULTS. Head: Vertex with decumbent scales silver or dark, broad, narrow
curved or narrow spatulate; scales along median longitudinal line in most species
silver, narrow curved and more lateral scales dark, broad; anterior patch of porrect
scales absent; occiput with erect scales dark or silver, usually scales of 1 color pre-
dominating; palpus entirely dark scaled; torus without scales. Thorax: Integument
dark brown to black; scales dark or silver, rarely a few golden scales present; meso-
notal disc with narrow curved scales, usually not transversely silvered in females,
often transversely silvered in males, the silvering usually an anterior band, less fre-
quently most of or entire disc silvered; complete silver acrostichal and posterior
dorsocentral lines present or absent, often present only in males; acrostichal setae
present or absent; fossal and supraalar areas each with variably developed silver mac-
ula, the maculae in females joined or disjunct, in males always joined; prescutellar
area with or without marginal silver scales, central portion without scales; lobes of
scutellum with or without silver scales; apn, ppn, ppl, paratergite, pra, upper stp,
posterior stp and mep with patch of scales, ssp with or without scales; all pleural
scales silver (rarely dark on ppn); apn, ppl, posterior ppn, psp, pra and posterior stp
and upper mep with setae. Legs: Midtarsi and hindtarsi usually with broader silver
bands than foretarsi; femur I usually with ventral or posterior patch of silver scales
and usually without knee spot, when present a single row of apical silver scales; tib-
ia I usually entirely dark, rarely with posterior silver streak; tarsus 1-I with basal
complete or incomplete silver band, with or without apical silvering, when present
usually at most a narrow band; 2-I with or without basal silvering, when present
usually at most a narrow band; 5-I rarely silvered; tarsus I otherwise dark; femur
II usually with ventral or posterior silver line and knee spot; tibia II dark; tarsus
1-II usually with basal and apical silver band, sometimes entirely silvered, infre-
quently only narrow basal band present; 2-II usually with basal silver band, some-
times most or all of segment silvered; 5-II infrequently silvered; tarsus II otherwise
dark; femur III usually with dark basal band or at least some dark basal scales, al-
ways with dark subapical band, remaining portion of segment silvered; tibia III dark;
tarsus 1-III usually with basal and apical silver band; 2-III with basal silver band;
S-III infrequently silvered; tarsus III otherwise usually dark (in occasional specimens
24 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
apex of 2-III and base of 3-III with a few silver scales). Wing: Silver scales usually
present at base of veins C and R and sometimes at base of Cu, often forming lines
reaching crossvein h (or the level of A in case of vein R); wing scales otherwise
dark. Abdomen: Laterotergite silver scaled; tergites II-VIII in females and II-VI
in males with lateral patch of silver scales, VIII in males usually entirely silver scaled;
sternites II-VII with basal silver band. Female Genitalia: Cercus short and broad,
segment VIII only slightly retracted.
MALE GENITALIA. Sidepiece: Basal tergomesal area with or without dense tuft
of setae: median sternomesal area with or without sclerite produced into mesal
membrane, sclerite sometimes with tuft of setae. Prosophallus: Filament typically
stout, setalike and with terminal hook, usually without structural elaborations but
in argyrothorax distally expanded into foliate structure. Aedeagus: Basal portion
subequal in diameter to or broader than distal portion; truncate distally.
PUPAE. Cephalothorax: Trumpets much darker than integument of cephalotho-
rax. Abdomen: Hairs 9-II-V and usually 9-VI caudad of 6-II-VI; hairs 10-VI much
closer to 7-VI than to each other. Paddle: Apex produced, evenly rounded or weak-
ly emarginate.
LARVAE. Head: Hair 4-C inconspicuous, multiple, usually caudad of 6-C, some-
times slightly cephalad of 6-C; hair 5-C well caudad of 7-C; 6-C slightly cephalad
of 7-C; hair 5-C interdistance subequal to that of 6-C; hair 15-C subequal in length
to mental plate; mental plate triangular in outline, the teeth more or less similar in
shape but broader laterad. Antenna: Hair 1-A single to 4-branched. Thorax: Hairs
1-3-P on common tubercle; 5-7-P on separate tubercles, only tubercle of 7-P well
developed; 6-P situated about midway between 5,7-P. Abdomen: Hair 12-I present.
Segment VIII: Comb scales 19 to more than 100, in 2 or more rows; free portion
with uniform fringe of very fine spicules at apex or along entire margin; apical scales
with free portion usually spatulate to pandurate, rarely awl-shaped. Siphon: More
or less evenly tapered; length 0.53-1.07 mm; classical index about 2.5-3.3; L/S 1.82-
2.86; P/L 0.44-0.64; H/L 0.51-0.69; pecten teeth with basal denticles, distal teeth
not detached. Anal Segment: Saddle finely spiculose, the posterior dorsal spicules
longer but not developed as conspicuous spines; hair 1-X inserted on saddle; 3-X
single; ventral brush without boss, with 10-16 hairs; 4a-X greater than half length
of longest hair of brush.
DISCUSSION. The above descriptions include primarily features of the Terrens
Group that separate it from other groups of the New World Finlaya. The system-
atics and evolution, the distribution, the bionomics and the disease relations of the
group are discussed in the preceding chapters.
KEYS TO SPECIES
FEMALES
(5. braziliensis and 15. impostor unknown)
E Supraalar macula broadly joined to fossal macula (fig. 17) except in some
insolitus and alboapicus; femur III with basal dark band complete and
broad, at least 0.09 or mesonotal disc transversely silvered; pra hairs
often dark, more heavily pigmented than those of pp/ and usually than
those of upper mep; vein C often with basal line of silver scales (devel-
oped on anterior and/or dorsal surface) reaching crossveinh . . . .2
2(1).
3(2).
4(3).
5(4).
6(S).
7(5).
8(7).
9(8).
10(Q9).
Schick: Terrens Group of Aedes (Finlaya) 3 25
Supraalar macula usually not joined to fossal macula; femur III with bas-
al dark band usually incomplete, if complete less than 0.09; mesonotal
disc not transversely silvered; pra hairs pale (except in some galindoi);
vein C without long basal line of silver scales as above, at most with
linée-rarely: reacminmO SD: torctossven Wich Box a iene Bae 4S
Ppn with scales dark; fossal macula with some pale gold scales in addition
to the silver (Aitkeent Subgroup) . . > .ohewl 13: artkeni
Ppn with scales silver; fossal macula with all scales silver 7) DEL AS
Tarsi 1-I], HI without apical silver band and tarsi 2-II,III without basal sil-
ver band; upper stp and mep scale patches contiguous (fig. 25) (Buena-
ventura Subgroup) Enos . . . .10. buenaventura
Tarsi 1-II,[JI with apical sibven band. and ens 2-I1,UI1 with basal silver
band; upper stp and mep scale patches not contiguous (fig. 7). . . .4
Tarsus 5-III silver scaled; supraalar macula often not joined to fossal mac-
ula (Alboapicus Subsroup) . . 9. alboapicus
Tarsus 5-III dark scaled; supraalar macula broadly joined to fossal macula
oS
Vertex with all decumbent scales broad (Thorntoni Subgroup) . . .6
Vertex with decumbent scales narrow curved at least along longitudinal
midline.
Mesonotal disc with anterior silver band (fig. — vein R without silver
BOGIES. ye . . . 1. thorntoni
Mesonotal disc not transversely silvered (fie. 12), vein R with small basal
patch ofisilver:scalese! uo) Jeyiescius yo. cae his als - Aecargyrothorax
Mesonotal disc with broad anterior silver band (fig. 11); femur III with
basal dark band incomplete (Bertrami Subgroup) . . . . 3. bertrami
Mesonotal disc not transversely silvered; femur III with basal dark band
Comstlete xii: lecigacdiy, sensing 3. deed mathe tee utes Temes
Occiput with all erect scales dark; ssp scales absent; vertex with decum-
bent scales along median longitudinal line generally dark and with scales
laterad of line narrow curved acaba Subgroup)
sk metoecopus
Occiput with all erect scales pale or - with mixture of pale and dark scales;
ssp scales usually present, when absent vertex with median longitudinal
line of silver scales and with scales laterad of line broad (Terrens, In-
solitus Subgroups)
Ssp scales absent... 46. ede we el ee ee 2 eee ee
SP SOROS TP ORRIN ii eae ek Saat a ile 9 Baas 6 02 a kine ea
Pra hairs usually pale; fossal macula usually broadly reduced mesad and
often reduced laterad (figs. 15,29); acrostichal setae present or absent
1]
26
11(10).
12(10).
13(1).
14(13).
15(14).
16(15).
17(13).
18(17).
Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
Pra hairs dark, more heavily pigmented than those of ppl; fossal macula
narrowly reduced mesad and always reaching lateral margin of fossa
(fies Tilo} acrostichal setaeabsenis sete a eG Oke peed
Fossal macula a well delineated marking, often in form of sublateral line,
sometimes divided into anterior and posterior segments (figs. 7,15);
mep with scale cee undivided or divided into dorsal and ventral por-
Hons... . . .4, terrens
Fossal macula usually a poorly defined and irregularly reduced marking
(fig. 29); mep with scale patch not divided . . . . . . 12. insolitus
Upper mep hairs pale, less heavily pigmented than those of pra; vertex
with decumbent scales laterad of median ie cele line narrow cur-
Wed . . 6. zavortinki
Upper mep hairs dark, as - heavily pigmented as those of pra; vertex with
decumbent scales laterad of median longitudinal line broad
es apollo
Acrostichal setae absent; tarsus 1-II] with median dark band usually in-
complete or complete and narrow, at most about0.33 .... . 14
Acrostichal setae present; tarsus 1-II with median dark band usually -com-
pleteand broad; 0.33 orgreater 2) Woe ee Cee ee ee
Tarsus 5-III silver scaled; tarsus 1-I with broad apical silver band, about
0.4 (Diazi Subgroup) . . y eee 28. diazi
Tarsus 5-II] dark scaled; tarsus 1-1 at most with very narrow apical silver
Pea rina ah neem en Tag an Se OER re ges yee eee i ae
Proboscis shorter than or subequal in length to femur I; femur II with
knee spot moderately broad, the silver scales at most just reaching an-
terior subapical setae (Podographicus Subgroup) .
Ci2S ‘podographicus
Proboscis longer than femur I: femur 1 with Knee spot broad, the silver
scales extending basad of anterior subapical setae (Tehuantepec Sub-
Ps ogee eee ee rem e Pea a Fei Sato Sh
Vertex with all decumbent scales silver . . . Pe emi 0 fe tehuantepec
Vertex with silver and dark decumbent scales, latter forming submedian
Pe eee aay ge at ee Se Sg BSR Up 8% le eee ROCIO
Femur II with or without knee spot, when present, narrow, a single row of
apical scales; ssp scales absent (Galindoi Subgroup). ..... . 18
Femur II with broad knee spot, the scales extending basad of anterior sub-
apical setae; ssp scales present or absent (Homoeopus, Heteropus Sub-
RUE eas craig ao aye cg ged ag Pcs NR a Ba AN IE
Fossal macula reduced only mesally (figs. 48,50); supraalar macula joined
to fossal macula; femora I,II with well developed posterior patch of sil-
Wermsesietey hp re AR ee i ae GA are et a 2 aryl
19(17).
20(19).
21(20).
22(20).
zoka2).
24(23).
yp
Schick: Terrens Group of Aedes (Finlaya) 27
Fossal macula reduced anteriorly and mesally (fig. 44); supraalar macula
disjunct from fossal macula; femora I,II without posterior patch of sil-
ver scales 2. i har Pe At. Se Sogalindeis 22: campana
Ssp scale patch absent although 1 or 2 scales sometimes present .
RR CACORE SRN GT AE i 7d | eae oe CLEMO S81 3 . 20. vargasi
SSD Stale Pare Pree oe a ee Glee ate ni
Proboscis shorter than or subequal in length to femur I; midlobe of scutel-
lum with silver scales . . . Se aca
Proboscis usually longer than, sometimes subequal in n length to femur I,
when subequal midlobe of scutellum without silver scales . . . . 22
Acrostichal line absent or represented by scattered silver scales (fig. 39);
midlobe of scutellum usually with a mixture of silver and dark scales,
the dark scales usually predominating, infrequently all scales silver .
. . 19. sumidero
Acrostichal line present, complete (fig. 35): midlobe of scutellum with
all scalesigilver (> oF gin Coe Sertina ie Gay Gon tie = ee
Femur I with narrow knee spot; complete and strong acrostichal and pos-
terior dorsocentral lines present (fig.37). . . . . . . .16. amabilis
Femur I without knee spot; complete and strong acrostichal and posterior
dorsocentral lines usually absent, sometimes complete but weak. . 23
Vertex with decumbent scales laterad of median longitudinal line broad
. . . . 18. idanus
Vertex with decumbent scales laterad of median longitudinal line nar-
LOW CHEV OGS. See ae a eae ee ee ah
Midlobe of scutellum without silver scales; proboscis subequal in length to
orlongerthanfemurlI. . » Go vey ib2yheteropus
Midlobe of scutellum usually with silver. scales: proboscis longer than fe-
ee U5 ata a, Se TR os ee ee ote Bomoecapns
MALES
(3. bertrami, 8. berlini, 12. aitkeni, 16. amabilis,
27. schroederi and 28. diazi unknown)
Vein C with basal line of silver scales reaching crossvein h; mesonotal disc
usually transversely silvered. . . “2
Vein C with small basal patch of silver scales or line ‘teaching at most to
about 0.5 to crossvein h; mesonotal disc not transversely silvered . 12
Vein R with basal line of silver scales much longer than that of vein C (Ho-
moeopus Subgroup). . . an
Vein R with basal line of silver ee fick ene thank that ce vein ne
. 4
28
3(2):
4(2).
5(4).
6(4).
7(6).
8(6).
9(8).
10(9).
Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
Median sternomesal area of sidepiece with well developed tuft and usually
with well developed convexity and sclerite (fig.31). . 14. homoeopus
Median sternomesal area with these structures not strongly differentiated
(ie SO ee Sa ee ts oS: impestor
Ssp scale patch absent . ee ee re ee es Pe
Ssp Scale paten presenti: a i ee eee ork ee
Tarsi 1-II,III with apical silver band and 2-II,III with basal silver band;
tarsus 5-III silvered; upper stp and mep scale patches not contiguous
(fig. 7) (Alboapicus Subgroup). 8 . . . 9, alboapicus
Tarsi of legs II,III without above silver markings; upper stp and mep scale
patches contiguous (fig. 25) (Buenaventura Subgroup)
Si Dee A ais Sega 10. buenavéitura
Vertex with scales along longitudinal midline broad (Thorntoni Subgroup)
Vertex with scales along longitudinal midline narrow curved .... .8
Mesonotal disc with anterior silver band not emarginate posteriorly (fig.
9); palpus 1-2 labellum lengths shorter than proboscis. Claspette fila-
ment not expanded distally (figs.9,11) . . . . . . . .1. thorntoni
Mesonotal disc with anterior silver band emarginate posteriorly (fig. 13);
palpus 4-7 labellum lengths shorter than Leeda Claspette filament
expanded distatby(ige. 13). oe . . . 2, argyrothorax
Mesonotal disc transversely silvered for most its length (except in Trinidad
where silvering sometimes restricted to anterior half of disc or trans-
verse silvering not developed), silvered area not distinctly emarginate
posteriorly (fig. 29); vertex with decumbent scales laterad of median
longitudinal line silver, broad (Insolitus Subgroup) . . . .12. insolitus
Mesonotal disc not transversely silvered or with transverse silvering devel-
oped as anterior silver band less than half length of disc and deeply
emarginate posteriorly (fig. 17); vertex with decumbent scales laterad
of median longitudinal line dark or silver, narrow or broad (Terrens
COULD) STCCAUPO) AGS TARE SSeS SOs SA ec ee Ue
Mesonotal disc without or with very narrow and barely discernable anter-
ior band (fig. 15); pra hairs pale; vertex with decumbent scales laterad
of median longitudinal line narrow curved (Brazil) . . . . .4. terrens
Mesonotal disc with relatively broad anterior silver band (fig. 17); pra hairs
dark, more heavily pigmented than those of ppl; vertex with decum-
bent scales laterad of median longitudinal line narrow curved or broad
10
Hairs of pra and upper mep about equally dark; vertex with decumbent
scales laterad of median longitudinal line broad (Colombia). . 7. apollo
Hairs of pra more heavily pigmented than those of upper mep; vertex with
decumbent scales laterad of median longitudinal line narrow curved
KS) CE CMa Sg RRCCRNE Heecal ek IG Mamet t uy eh 0) NO aha i OO ee |
11(10).
12(1).
13(12).
14(13).
15(12).
16(15).
17(16).
18(15).
19(18).
Schick: Terrens Group of Aedes (Finlaya) 29
Vertex with decumbent scales laterad of median longitudinal line broad |
(French Guiana, Brazil) . . . . 3. braziliensis
Vertex with decumbent scales laterad of median longitudinal line nar-
row-curved(Panania) ii. i trea. Gea Ola eS Gy Zavortinks
Acrostichal:setae absent rn 4 a ee eee ce BS
Acrostichal Seiae Dreseme iia a ue Br a ae ae Ot nS
Occiput with erect scales dark (Metoecopus Subgroup) . 11. metoecopus
Occiput with érect Seales pale fo: Oa ep Baie BG) Aone ee eee
Palpal segment 3 without prominent tuft of setae, the setae at ventrolater-
al apex shorter than segments 4 and 5 combined. Prosophallus with lat-
eral portion of mesal lobe usually moderately inclined, between 15°
and 30° from horizontal; stems usually bowed and convergent (figs.
52,54,56) (Podographicus Subgroup) . . . . . . 25. podographicus
Palpal segment 3 with prominent tuft of setae as long as segments 4 and 5
combined. Prosophallus with lateral portion of mesal lobe slightly in-
clined, about 15° or less from horizontal; stems not bowed, divergent
or essentially oie pes aes (Tehuantepec sat eae
ave ; nas tehuantepec
Femur II without or with narrow knee spot, the silver scales a single row
at apex of segment; ssp scale patch absent (Galindoi Subgroup) . . 16
Femur II with broad knee spot, the silver scales extending basad of anter-
ior subapical setae; ssp scale patch present (Heteropus Subgroup) . 18
Femora I,II with well developed posterior patch of silver scales. Sidepiece
with median sternomesal tuft poorly differentiated and the setae not
wavy (figs.48,50) .... . . . . 24, daryi
Femora I,II without posterior patch of silver scales. Sidepiece with median
sternomesal tuft well differentiated and the setae wavy (figs. 44,46) .
Prosophallus with median lobe projecting farther cephalad than lateral
lobe (fig. 44) .... «ay big, wey MORIN!
Prosophallus with median lobe projecting to about same level as lateral
loDe (MEAG) 5a Pag ea a Bee eanmaena
Median sternomesal area of sidepiece with sclerite and tuft well developed;
hook of filament strongly angulate (figs.35,37) . . . .... . 19
Median sternomesal area of sidepiece with sclerite and tuft absent or poor-
ly developed; hook of filament not strongly angulate (figs. 39,40,42) .
Palpus subequal in length to or slightly longer than proboscis. Basal tergo-
mesal area of sidepiece without dense patch of long setae (fig. 35) .
17. gabriel
Palpus about 2 labellum lengths shorter than ‘proboscis. Basal tergomesal
area of sidepiece with dense patch of long setae (fig. 37). . 18. idanus
30
20(18).
21(20).
Zt).
Sty.
4(3).
5(4).
6(4).
7(6).
Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
Ssp scale patch absent; mesonotum without complete acrostichal or post-
erior dorsocentral lines (fig. 40); tarsus 1-II with median dark band in-
COMP tees ; . . 20. vargasi
Ssp scale patch eeceenk: mesonotum with complete acrostichal or poster-
ior dorsocentral lines; tarsus 1-II with median dark band complete, a-
DOO enh eo ae ote Ot al
Acrostichal line absent; posterior dorsocentral line complete (fig. 39);
segment 3 of palpus with apical ventrolateral tuft not as long as seg-
ments4and5 combined. . . . . 19. sumidero
Acrostichal line present, complete, sometimes weakly developed: posterior
dorsocentral line incomplete (fig. 42); segment 3 of palpus with apical
ventrolateral tuft as long as segments 4 and 5 combined .
wk, heteropus
LARVAE
(3. bertrami, 5. braziliensis, 16. amabilis,
and 19. sumidero unknown)
Hair 5-C usually with 4 or more branches (when fewer branches, only on 1
side) and/or 14-P branched . ... . ee
Hair 5-C usually single or double, sometimes triple (when | more ‘branches,
only on | side) and 14-P usually single (when rarely branched, only on
eg an cons awe Bact Atroy 1) spat ethene
Hair 5-VII cephalad of 4-VIT . . . . . . . +. . . .10. buenaventura
Eight ory he Cae Oa ee par he te NER TOA AGE RNR. ri See eg
Hairs 4-VII and 3-VI branched. . . . . . 9, alboapicus
Hairs 4-VII single and 3-VI a single (when rarely branched, only on
Oriosecr ee er ee
Hair 2-II well mesad of 4-II (fig. 42);14-P single. . . a
Hair 2-II mesad of 4-II for about only 1 alveolus width, often laterad of
at (ee Se baer rane te ee ae
Hair 11-P less than half length of 14-P; 1-VIII shorter than 2-VIII; 6-C
single or double; bmh single. . . . . . 18. idanus
Hair 11-P about half length of 14-P; 1-VIII usually at least subequal in
length to 2-VIII; 6-C usually with more than 2 branches; bmh usually
branched but often ee re a ee te 21. heteropus
Hair 14-C usually with 3 or more branches; 1-A usually branched but of-
ten single; free portion of apical scales ligulate, awl shaped or spatulate
. . . 24, daryi
Hair 14. ‘usually with fae rece 3 ‘branches: af ie usually single (when
branched, only on 1 side); free portion of apical scales spatulate. . .7
Comb scales 35-47, in 4 rows; free portion of midapical scale longer than
SORSIS BOSON er eG eee a aa
8(1).
9(8).
10(8).
11(10).
12(11).
£312).
14(13).
15(11).
16(15).
Schick: Terrens Group of Aedes (Finlaya) 31
Comb scales 23-32, in 3 rows; free portion of midapical scale shorter than
or subequal in length to sessile portion. . . . . . . . 23. campana
Hair 7-C short, less than half length of 6-C; 11-C short, less than length of
Herb £50 y Ais D
Hair 7-C more than half length of 6-C: 1 1c longer than mentum. nee
Hair 8-S single; 2-A about 2.0 distal portion of 6-A; anal saddle extending
less than halfway around segment (fig. 14) . . . . . 2. argyrothorax
Hair 8-S multiple; 2-A about 3.0 or more length of distal portion of 6-A;
anal saddle extending more than halfway around segment (figs. 10,12)
.1. thorntoni
Hairs 14-Cand bmh usually branched, rarely single; 11-P usually at least
half length of 14-P; 6-C usually double ortriple . . . . 12. insolitus
Hairs 14-C and bmh usually single, rarely branched; 11-P less than half
length of 14-P; 6-C usually single, sometimes double, rarely triple .
1]
Hair 4-VII at least double; 4-M and 3-III usually at least triple; 10,12-VII
often branched. . . yi 2 eeeae
Hair 4-VII usually single’ rately double; 4- M and es Il with less than 3
branches: Tel 7 ey FH sinelewar 4, arity to ge el gre bey cee! 1S
Ventral brush with 11 or 12 hairs; 4a-X with 6 branches; 10,12-VII single
. 8. berlini
Ventral brush with 3 or more 2 hairs: 4a- X usually with 8 or more branch-
es; 10,12-VII usually double but sometimes single . . . . . . . 13
Hair 2-VIII usually branched; 6-C often branched but usually single
get Fe EC Ae Uae aati nese is hol ei See
Hairs.2-V hand GAC betipsingie- i: cit ie eo? ed eee ee ee A
Hair 7-C with 5-9 branches 5-C usually branched, sometimes single
Wt She RIS (nS Sal ea ia ce eae ea Ais 2 ee ee nee 6. zavortinki
Haiti." tripe; Gar simele oy. peepee Riemaes Beis nee eeo
Saddle extending around segment to at most moderate distance beyond
horizontal midline, submarginal slit absent, ventral margin either with
broad rounded incision or irregular in outline (fig. 52) ;
bee Rete metoecopus; 14. homoeopus;
16. impostor; 17. gabriel; 20. vargasi; 26. podographicus
Saddle extending around segment far beyond horizontal midline, with
ventral submarginal or marginal slit (figs.59,60). ...... . #16
Comb scales 33-54, in 4 rows, narrow (fig.59) . . . . 27. tehuantepec
Comb scales 20-29, in 2-3 rows, stout (fig.60) . . . . .28. schroederi
32 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
Thorntoni Subgroup
ADULTS. Head: Vertex of both sexes with decumbent scales along longitudinal
midline broad; occiput of both sexes with all erect scales pale; proboscis of females
longer than femur I; palpal segment 3 of males usually with 1 or 2 short apical ven-
trolateral hairs; segment 4 with hairs of ventrolateral row usually short and sparse.
Thorax: Mesonotal disc of females with or without narrow anterior silver band, of
males with very broad anterior silver band; complete acrostichal or posterior dorso-
central lines absent in females; acrostichal setae absent; fossal macula of both sexes
coextensive with fossa; supraalar macula truncate posteriorly, in females broadly
joined to fossal macula; ppn silver scaled; ssp scales present; pra hairs of females
dark; upper stp and mep scale patches not contiguous. Legs: Mid- and hindlegs not
shaggy; mid- and hindtarsi with conspicuous silver markings; tarsus 1-I with or with-
out narrow apical silver band; femur II with knee spot broad, the scales extending
basad of anterior subapical setae; tarsus 1-I] with median dark band usually com-
plete and moderately broad; tarsus 2-II with complete apical dark band; tarsus 5-II
without silver scales; femur III with basal dark band complete, broad; tarsus 5-III
without silver scales. Wing: Vein C of females with basal line of silver scales reach-
ing about 0.5 to crossvein h or up to h, of males with line reaching h; vein R of
males with basal line of silver scales reaching about 0.5 to level of h; vein Cu of
males without silver scales.
MALE GENITALIA. Sidepiece: Basal tergomesal area with or without dense patch
of long setae; median sternomesal area without strongly developed sclerite; with or
without tuft; specialized subapical sternal seta absent. Prosophallus: Mesal lobe with
lateral lobe variously inclined but always 15° or less from horizontal; stem not bow-
ed; filament with hook not strongly angulate.
PUPAE. Cephalothorax: With or without pale inverted V-shaped marking; hair
SC less than twice length of 4C; 9-C single. Abdomen: Hair 1-I with primary
branches predominantly multiple; 2-II mesad of 3-II for at least 0.2, usually 0.3 or
more the distance from 1-II to 3-II; 3-III usualiy branched. Paddle: Clear; apex not
produced; hair 1-P shorter than paddle.
LARVAE. Head: Hairs 5,6-C single; 7-C less than half length of 6-C; 11-C shorter
than mental plate; 14-C shorter than or subequal in length to mental plate, single;
bmh single. Antenna: Hair 1-A single. Thorax: Hair 11-P less than half length of
14-P; 14-P single; 3-M branched; 4-M double to multiple; 8-T less than half length of
metathoracic pleural tubercle. Abdomen: Hair 5-I shorter than 4-I; 2-II well mesad
of 4-II; 3-III double to multiple; 10-III, 3-VI and 4-VII branched; 5-VII caudad of 4-
VII, less than half length of 3-VII; 10,12-VII branched. Segment VIII: Midapical
comb scale with free portion shorter than or subequal in length to sessile portion;
hair 2-VIIJ branched. Anal Segment: Saddle extending around segment for short to
moderate distance, ventral margin withoui deep slit.
DISCUSSION. The Thorntoni Subgroup shows several characters which are unique
in the Terrens Group. In the adults the decumbent scales along the longitudinal mid-
line of the vertex are broad, in the pupae hair 3-III is usually branched, and in the
larvae hairs 7,11-C, 8-T and 5-I are very short and hair 2-VIII is often triple to 4-
branched (the latter hair being single or double in the other species). Other charac-
ters shown only infrequently among the other subgroups are the truncate posterior
margin of the supraalar macula, the broad anterior silver band of the male mesono-
tum and the clear pupal paddle.
The subgroup shows all of the features that characterize the Terrens phyletic
Schick: Terrens Group of Aedes (Finlaya) 33
line and is presumed to have been derived from the more generalized Terrens Sub-
group (see chapter on Systematics and Evolution). It is distributed in the lowlands
of Central America, principally on the Atlantic side, and in the Atlantic drainages
of South America as far south as Rio de Janeiro.
The 2 species of the subgroup, thorntoni and argyrothorax, are evidently closely
related since they share the singular subgroup characters mentioned above. The male
genitalia of these species, however, are strikingly different, those of argyrothorax
being unique in the Terrens Group. It would seem that argyrothorax has been de-
rived from a thorntoni-like ancestor.
1. Aedes (Finlaya) thorntoni Dyar & Knab
Figs. 2,9-12
1907. Aedes thorntoni Dyar and Knab, 1907:10. TYPE: Holotype ?, Bluefields, Nicaragua, W.
F. Thornton [USNM, 10143]. Type data from Stone and Knight (1956:225).
Aedes (Finlaya) thorntoni of Dyar (1921:153; 1925b:147-148; 1928:225-226); Bonne-Wepster
and Bonne (1925:361,365,419-420); Costa Lima (1930:257-258); Edwards (1932:150); Lane
(1936a:11); Knight and Marks (1952:549); Horsfall (1955:462).
Aedes thorntoni of Theobald (1910:485); Howard, Dyar and Knab (1917:819-821); Dyar (1918:
73,80).
Aedes (Finlaya) terrens in part of Arnett (1949:227); Lane (1939:105; 1953:686-687); Stone,
Knight and Starcke (1959:171); Belkin, Schick and Heinemann (1965:395).
Aedes terrens in part of Kumm, Komp and Ruiz (1940:416,450).
Aedes insolitus in part of Busck (1908:64).
FEMALE. Head: Vertex with all decumbent scales broad, most or all dark. Thor-
ax (fig. 9): Mesonotal disc typically with anterior silver band about 0.33-0.75 length
of fossa; short silver acrostichal line often projecting caudad from anterior silver
band; prescutellar space and scutellum without silver scales. Legs: Femur I with
small posterior patch of silver scales in basal half; femur II without or with only a
few posterior silver scales; tarsus 1-I] with median dark band about 0.33-0.67; femur
III with basal dark band 0.13-0.19 (0.11-0.21), subapical dark band 0.18-0.23 (0.16-
0.23); tarsus 1-III with basal silver band 0.12-0.17 (0.09-0.19), apical silver band
0.17-0.21 (0.15-0.23). Wing: Vein C with basal line of silver scales reaching crossvein
h; vein R without silver scales.
MALE. Head: Palpus usually about 2 labellum lengths shorter than proboscis,
infrequently 1 labellum length; segment 3 usually with 1 or 2 short apical ven-
trolateral hairs and segment 4 with hairs of ventrolateral row usually short and
sparse, but in Portobello area of Panama segment 3 with apical ventrolateral tuft of
hairs as long as segments 4 and 5 combined and segment 4 with ventrolateral row of
long, closely spaced hairs. Thorax (fig. 9): Mesonotal disc with about anterior 0.67
transversely silvered, posterior margin of silvered area usually an almost straight line;
acrostichal and dorsocentral lines,often projecting caudad from transverse silvered
area; pra hairs pale or dark. Legs: Tarsus 1-II with median dark band complete,
about 0.25-0.67.
MALE GENITALIA (figs. 9,11). Sidepiece: Length 0.25-0.30 mm: without spec-
lalized tufts of setae. Prosophallus: Length 0.07-0.10 mm; width 0.11-0.12 mm
(0.10-0.12 mm); mesal lobe with lateral portion usually not appreciably inclined
34 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
or declivous mesad, usually inclined less than 15° (at about 15° in 1 specimen);
stems parallel when short (fig. 11), divergent when long (fig. 9); filament not ex-
panded distally, ratio 0.60-1.30. Aedeagus: Length 0.10-0.12 mm.
PUPA (figs. 9,11). Abdomen: Hair 2-II mesad of 3-II for 0.3 or more the distance
from 1-II to 3-II, usually at least 0.4; 3-III usually branched [in most localities
branched in both sexes; in Portobello area, Panama, single in male, branched in fe-
male].
LARVA (figs. 10,12). Head: Hair 7-C single to 6-branched [in most localities usu-
ally triple or 4-branched, in Portobello area usually double or triple]. Antenna: Hair
1-A reaching apex of antennal shaft; 2-A at least 3.0 length of distal portion of 6-A.
Thorax: Hair 1-P usually double or triple (1-3); 4-P usually double or triple (2-4); 5,
7-P usually single or double (1-3); 3-M usually triple to 4-branched (2-4); 4-M usually
5-6 branched (4-6). Abdomen: Hair 3-III usually 5-6-branched (3-7); 10-III double
to 4-branched; 3-VI double to triple (1-3); 4-VII usually triple to 4-branched (3-6);
10-VII double or triple; 12-VII usually triple or 4-branched (2-4). Segment VII:
Comb scales about 75 to more than 100; free portion of midapical scale shorter than
or subequal in length to sessile portion, usually shorter, length 0.020-0.024 mm
(0.018-0.027); hair 2-VIII usually triple to 4-branched (2-4). Siphon: Length 0.61-
0.87 mm; L/S 2.63-2.78 (2.57-2.86); P/L 0.47-0.52 (0.46-0.54); H/L 0.55-0.59
(0.54-0.60); hair 8-S branched. Anal Segment: Saddle extending more than half-
way around segment; ventral brush usually with 12 hairs (11-13); 4a-X usually 10-
branched (9-14).
SYSTEMATICS. Aedes thorntoni can be differentiated from all the other species
of the Terrens Group in the adults by the form of the silver markings of both sexes —
and in the larva by the often highly branched condition of hairs 3,4-M, 3,10-III,
4,10,12-VII and 2-VIII.
The population of the Portobello region of Panama differs from the others in 2
obvious respects, (1) the male palpus shows a denser and longer vestiture of hairs
and (2) pupal hair 3-III is single in the male and branched in the female (6 speci-
mens) whereas in the other populations the hair is branched in both sexes.
DISTRIBUTION (fig. 2). Atlantic lowlands, from Nicaragua to Panama; Canal
Zone; Pacific lowlands of provinces of Panama and Darien, Panama. Material ex-
amined: 803 specimens; 240 6, 229 2, 185 pupae, 148 larvae; 103 individual rear-
ings (75 larval, 25 pupal, 3 incomplete). For the Komp collections see the explana-
tory chapter.
COSTA RICA. Limon: Chase [elev. 0-700 ft] , 1 ¢ (221) [USNM].
NICARAGUA. Zelaya: Bluefields, W. Thornton, 6 ? (10), type series [USNM].
PANAMA. Bocas del Toro: Bocas-Chiriqui road, nearest town Almirante (elev. near sea level), 6
May 1943, treehole, 1 pd (PA 328-103) [UCLA]. Canal Zone: Barro Colorado Island [elev. 100-
500 ft], 24 July 1923, H. Dyar, R. Shannon, 1 ? (P29-II); 27 July 1923, H. Dyar, R. Shannon, 1
2 (P29-3); 28 July 1923, H. Dyar, R. Shannon, 1 pd (P29-1); 16 Aug 1923, H. Dyar, R. Shannon,
1 2 (P29-II); Jan 1935, 1 6 [USNM] ; 26 July 1935, L. Rozeboom,3 ¢ (PAR 81) [UCLA]; 15 Apr
1939, W. Komp, 1 ? (207E-3); 8 Jan 1943, treehole, 1 d (207B-21); 7 May 1943, double treehole,
W. Komp, 1 1d (207A-2) — treehole, 1 1d (207A-33) — 1 6 (207B-34) — treehole, 1 lpé (207B-38);
21 May 1943, treehole, G. Fairchild, 1 Ipdé (207A-19); 21 May 1943, hole in stilt palm, W. Komp,
2 Id (207A-25 27), 4 19 (207A-5 26,40; 207D-20), 2 6, 2 1(207D-9); 21 May 1943, treehole, 1
1d (207A-39), 1 Ip? (207A-42) — small treehole, 1 1d (207D-15), 1 19 (207A-15) — 19,21(207A-
21) — 2 1 (207A-35) — treehole, 3 9, 2 1 (207A-38) — 3 9, 3 1 (207A-41) — treehole, 1
d, 3 2 (207A-44) — deep double treehole, 2 6, 1 9, 4 1 (207B-7) — large treehole, 1 19 (207B-11)
— stilt roots, 1 d, 1 9 (207D-12); May 1943, treeholes, W. Komp, 2 6 (207A-32); 26 April 1945,
treehole, W. Komp, 1 12 (207E-15); 15 May 1945, treehole, W. Komp, 1 lpé (207C-21; 207D-3), 4
Schick: Terrens Group of Aedes (Finlaya) 35
Ip? (207C-29; 207E-16,19,30), 4 6, 4 ? (207C-27), 1 6, 3 2 (207C-43), 2 %, 2 1 (207D-30), 1 1
(207C-15) — treehole, 2 lp? (207C-40; 207D-37) — treehole, 1 6, 1 P, 1 p, 2 1 (207C-44), 14,1
2, 2 p, 3 1 (207D-24) — 1 lpdé (207C-4) — treehole, 1 6 (207C-18) — 1 lpd (207C-31) — treehole,
1 lpd (207E-31) — 1 1d (207E-34); 22 May 1945, large treehole, W. Komp, 3 Ip? (207C-30,36;
207D-44), 4 6 (207C-37), 1 6 (207C-32), 1 6 (207D-33), 2 6, 1 2 (207D-38), 1 2 (207E-23)
— treehole, 1 12 (207D-35), 1 6, 1 p, 21(207E-25), 2 6, 5 2? (207D-34), 2 9, 21(207C-9) — 1 Ip?
(207C-16) — 1 6 (207C-39) — low stump, 1 Ip? (207D-41); 23 May 1945, stilt palm, W. Komp, 1
Ipd (207D-42), 1 16 (207E-17), 4 6, 2 2 (207D-29) — 1 % (207D-25) — treehole, 1 6, 1 9, 2
1 (207E-14); 1945, treehole, W. Komp, 1 6 (207D-23) [UCLA; USNM] . Camacho, 14 Jan 1922, J.
Shropshire, 1 9; 17 Mar 1922, J. Shropshire, 2 9; 3 June 1922, J. Shropshire, 4 d, 1 °; 4 June
1922, J. Shropshire, 1 6; 8 Aug 1923, H. Dyar, R. Shannon, 1 9 (P64) [USNM] . Chiva Chiva, elev.
0-600 ft, Oct 1941, treehole, 1 ¢ (207B-42); 11 Nov 1965, treehole, height 2-4 ft, A. Quinonez
(PA 768-700,772,773), 7 lpé (769-20,22-24,26; 773-21 ,23), 7 lp? (769-21 27-29; 770-11; 772-10;
773-22), 6 pd (769-25 ,100; 770-100; 773-100-102), 3 IP (770-10; 772-11; 773-20), 2 9,5 p, 21
(768-4) 26 p, 2: L:(769-2),:2 3,12, 0-2, 1p, 210770-11 6,5 2S TL G72-1), 6.4,7-2,31p
(773-2) [UCLA] .Corozal [elev. 0-100 ft] , 13 Jan 1943, ‘‘ant treehole’’, 8 5, 16 2 (207B-22); 6 May
1943, 1 6, 2 2 (207B-36), 1 ¢ (207B-20); 12 Jan 1944, bamboo, W. Komp, 1 * (207D-17), 1 2, 2
1, 2 p (207D-14), 1 2 (207D-19); 3 May 1945, “‘square treehole”, W. Komp, 1 ¢ (207C-5) — tree-
hole, 1 Ipd (207C-17) — 1 6, 2 2 (207C-28); 4 May 1945, W. Komp, 1 16 (207C-12) — “‘square tree-
hole”, 1 6, 1 2 (207C-22) — treehole, 1 1d (207C-33); date not specified, 1 6 (207B-25) [UCLA;
USNM]. Corozal Damsite, 28 July 1943, bamboo, W. Komp, 1 6 (207A-36) — 1 6, 1 ? (207D-
18); Aug 1943, bamboo, W. Komp, 1 ¢ (207A-12); 18 Sept 1943, bamboo, W. Komp, 2 Ip?
(207A-3,33), 1 3 (207A-30); 12 Jan 1944, W. Komp, 2 6, 1 2, 3 1, 3 p (207D-16); 3 May 1945,
W. Komp, 1 d, 1 2, 2 1(207C-23) [UCLA; USNM] . Corozal Hospital grounds, 7 Sept 1943, treehole,
H. Herman, 1 Ip? (207A-16), 2 p? (207A-14,18) [UCLA; USNM]. Cruces Trail, Madden Forest,
elev. 200-600 ft, 17 Sept 1964, large treehole, height 5 ft (PA 707), 1 lpd (707-31), 2 lp? (707-36,
38), 1 p? (707-100), 7 L (707-3) [UCLA]. Empire [elev. 200-300 ft], 7 Aug 1923, H. Dyar, R.
Shannon, 1 6 (P56-1); 8 Aug 1923, H. Dyar, R. Shannon, 1 9 (P56-2) [USNM]; 7 Aug 1944, tree-
hole, Wood, Adams (ASM 86,88,89), 1 ¢ (86-2), 1 d (88-2), 3 6 (89-1) [UCLA]. Fort Randolph
[elev. near sea level] , 22 Apr 1922, J. Shropshire, 1 9 [USNM]. Fort San Lorenzo, 20 Aug 1923,
H. Dyar, R. Shannon, 1 6, 1 9 (P95) [USNM]. Fort Sherman [elev. near sea level] , 1916, L. Dunn,
13,1 9(C-53), 1 9 (C71=C53), 1 9 [USNM] ; 26 Oct 1948, 1 9 (25); 5 May 1949, 4 6 (174); 6 May
1949, 1 3; 12 Aug 1949, 3 gd; 19 Aug 1949, 1 36; 2 Sept 1949, 1 9; 9 Sept 1949, 2 6,1 9 [UCLA].
France Field [elev. near sea level], 1 Jan 1925, J. Shropshire [USNM]; 26 Oct 1948, 1 6 (35)
[UCLA]. Firjoles, elev. ca. 50 ft, 1 Dec 1965, large treehole, height 5 ft, R. Schick, A. Quinonez
(PA 844), 1 Ipd (844-10), 2 pd (844-101-102), 2 p? (844-100,103) [UCLA]. Gamboa [elev.
100-200 ft], 10 June 1943, Elton, 1 6, 1 9 (207D-21) [UCLA]. Gatun [elev. 100-200 ft],
8 Aug 1923, H. Dyar, R. Shannon, 1 6 (P61); 21 July 1926, D. Curry, 1 6, 1 9; 21 Aug
1926, D. Curry, 4 6, 3 2? [USNM]; 24 Sept 1964, treehole, height 15 ft (PA 712), 1 lpé
(712-34), 9 Ip? (712-30-33,35-39), 2 pd (712-104,111), 5 p? (712-101-103,112,114), 23 3, 13
?, 2 P, 54 p,6 L, 81(712-3) [UCLA]. Gold Hill [elev. 0-700 ft], 9 Dec 1921, J. Shropshire, 1
gd [USNM]. Juan Mina [elev. 100 ft], 18 Jan 1963, treehole, height 1 ft, 1 lp? (PA 5-101)
[UCLA]. Largo Remo [elev. near sea level], 27 July 1926, D. Curry, 3 6, 1 9, 11 Aug 1926,
D. Curry, 3 6, 1 @ [USNM]. Majagual, 3 Dec 1921, J. Shropshire, 9 6, 3 9; 14 Jan 1922, J.
Shropshire, 4 6, 3 9; 11 Feb 1922, J. Shropshire, 1 9; 22 Apr 1922,J. Shropshire, 2 ¢ [USNM].
Mandingo (? Mandinga), 13 Dec 1921, J. Shropshire, 4 g, 1 9; 14 Dec 1921, J. Shropshire, 1 3, 1
[USNM]. Margarita [elev. 0-100 ft], 21 Jan 1922, J. Shropshire, 3 6, 2 9; 12 Aug 1922, J. Shrop-
shire, 2 2; 19 Aug 1922, J. Shropshire, 1 6,3 9; 2 Dec 1922, J. Shropshire, 2 6, 1 9 [USNM]. Min-
di [elev. 0-100 ft], 28 July 1923, H. Dyar, R. Shannon, 1 6 (P40-II); 5 Aug 1923, H. Dyar, R.
Shannon, 1 9 (P40-1) [USNM]. Quarry Heights, 14 Sept 1949, S. Carpenter, 2 9; 15 Nov 1949, S.
Carpenter, 1 d [UCLA]. Rio Chagres, upper, treehole, A. Busck, 2 9 (141) [USNM]. Rio Curun-
du, upper, 23 Aug 1923, treehole, H. Crowell, 1 1d (207A-13) [UCLA; USNM]. Sweetwater [elev.
0-100 ft], 1 Mar 1922, J. Shropshire, 3 6,3 9 [USNM]. Summit [elev. 200-300 ft] , 13 Apr 1939,
W. Komp, 1 6 (207E-8); 13 Sept 1939, W. Komp, 2 ? (207E-2) — 1 9 (207E-6) — 1 2 (207E-9);
13 Dec 1939, treehole, W. Komp, 1 d (207E-4); 17 Aug 1941, high treehole in Mango, 2 ? (207B-
36 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
17), 1 9 (207B-16) — Ficus treehole, 2 ¢d (207B-31), 1 5 (207B-35), 1 2 (207B-26) [UCLA]. Ta-
bernilla [elev. 0-100 ft], 30 July 1908, A. Jennings, 2 ¢ (397,405), 1 ¢ (398.2); 14 Aug 1908, A.
Jennings, 4 2 (418,423 ,423.1,423.2); 22 Dec 1908, A. Jennings, 1 6 (458.2) [USNM]. Toro Point
[elev. near sea level] , 28 Jan 1922, J. Shropshire, 1 6; 13 Apr 1922, J. Shropshire, 4 6; 19 Apr
1922, J. Shropshire, 1 d [USNM]. Huile, near, elev. 100-300 ft, 7 Dec 1965, small treehole, height
4 ft, A. Quinonez (PA 875), 1 lp? (875-10), 1 L (875-1) (UCLA]. Locality not specified, 14 Feb
1924, C. Ludlow, 8 9; 25 Aug 1923, H. Dyar, R. Shannon, 1 6 (PII) .[USNM]. Colon: Portobello,
elev. near sea level, 5 Dec 1963, large treehole (PA 586), 1 lp? (586-102), 3 pd (586-101,103,105)
[UCLA]. Rio Caldera, elev. near sea level, 6 Dec 1963, large treehole (PA 592), 3 Ip? (592-101-
102,104), 1 IP (592-105), 8 L (592-2) [UCLA] . Darien: Jaque [elev. near sea level] , 3 July 1945,
W. Komp, 1 1; 4 July 1945, W. Komp, 6 1 [USNM]; July 1945, W. Komp, 1 6, 1 2? (207E-32),
1 d, 1 2 (207E-33), 1 6 (207E-35) [UCLA]. Santa Fe, elev. 200 ft, 22 Nov 1966, treehole, height
2-3 ft, O. Berlin (PA 944,945), 1 pd (945-100), 2 p? (944-100; 945-101); 9 Dec 1966, treehole,
height 15 ft, O. Berlin, Lineres (PA 992), 1 Ip? (992-10), 1 pd (992-101), 1 p? (992-100), 1 9
(992-100), 1 9 (992-1) [UCLA]. Panama: La Chorrera [elev. 0-300 ft], 17 Oct 1944, large tree-
hole, Adams, 1 6, 1 2 (ASM 210-1) [UCLA]. Panama [elev. near sea level], 9 Sept 1926, D.
Curry, 2 ¢ (26,26), 1 9 (25a) [USNM].
2. Aedes (Finlaya) argyrothorax Bonne-Wepster & Bonne
Figs. 2,13,14
1920. Aedes argyrothorax Bonne-Wepster and Bonne, 1920:179. TYPE: Holotype 3 (3925)
with genitalia (BB 353, M52), Geiersvlijt, an estate near Paramaribo, Surinam, near a tree-
hole [ITH]. Type data from Belkin (1968:4).
Aedes (Finlaya) argyrothorax of Dyar 1921:153; Bonne and Bonne-Wepster (1925 :422,424); Dyar
(1928:226); Shannon (1931a:8; 1931b:148); Edwards (1932:149); Chagas, Cunha et al (1937:
388); Kumm and Novis (1938:502); Lane (1939:101-102; 1953:688-690); Floch and Abonnenc
(1942a:5; 1942b:6; 1947:9-10,12); Cerqueira (1950:173-178); Del Ponte, Castro and Garcia
(1951:239-240); Knight and Marks (1952:523,546); Horsfall (1955:457); Stone, Knight and
Starcke (1959:159); Fauran (1961:28); Stone (1961:40); Belkin, Schick and Heinemann (1965:
63); Forattini (1965 :377 389-390); Belkin (1968:4).
Aedes (Gualteria) argyrothorax of Vargas (1950a:62).
Aedes oswaldi in part of Howard, Dyar and Knab (1917:819).
Gualteria oswaldi of Aiken (1909:12-13).
FEMALE. Head: Vertex with all decumbent scales broad, most or all dark. Thor-
ax (fig. 13): Mesonotal disc not transversely silvered, with a few scattered silver
scales along acrostichal line; prescutellar space with or without silver scales; scutel-
lum without silver scales. Legs: Femur I with well-developed posterior patch of sil-
ver scales in basal half; femur II usually without posterior patch of silver scales or
with patch represented by a few scattered scales, patch in Brazil sometimes well de-
veloped; tarsus 1-II with median dark band about 0.5; femur III with basal dark band
0.18-0.20 (0.16-0.21), subapical dark band 0.12-0.16 (0.11-0.17); tarsus 1-III with
basal silver band 0.08-0.10 (0.06-0.11), apical silver band 0.20-0.24 (0.19-0.27).
Wing: Vein C with line of silver scales extending about 0.5 to crossvein h or up to
crossvein itself; vein R with basal silver scales usually forming small patch, sometimes
a line extending about 0.5 to level of h.
MALE. Head: Palpus about 4-7 labellum lengths shorter than proboscis; segment
3 with 1 short apical ventrolateral hair; segment 4 with hairs of ventrolateral row
Schick: Terrens Group of Aedes (Finlaya) a7
short and sparse. Thorax (fig. 13): Mesonotal disc with about anterior 0.5 trans-
~ versely silvered, posterior border of silvered area shallowly emarginate; acrostichal
and posterior dorsocentral lines absent; pra hairs as in female. Legs: Tarsus 1-II
with median dark band usually complete, about 0.33-0.5, incomplete in some speci-
mens from the Guianas.
MALE GENITALIA (fig. 13). Sidepiece: Length 0.29-0.33 mm; basal tergomesal
area with dense tuft of fine setae; median sternomesal tuft present but not promin-
ent. Prosophallus: Length 0.13-0.14 mm; width 0.14-0.15 mm; mesal lobe with lat-
eral portion declivous laterad; stems convergent; filament foliaceous and striated (ra-
tio not calculated because of unique structure). Aedeagus: Length 0.15 mm.
PUPA (fig. 13). Abdomen: Hair 2-II mesad of 3-II for 0.2 or more the distance
from 1-II to 3-II, usually 0.3 or more; 3-III branched in both sexes.
LARVA (fig. 14). Head: Hair 7-C usually double or triple (2-4). Antenna: Hair 1-
A not reaching apex of antennal shaft; 2-A about 2.0 length of distal portion of 6-
A. Thorax: Hair 1-P usually single or double (1-3); 4-P single or double; 5,7-P single:
3,4-M double or triple. Abdomen: Hair 3-III usually double (2-3); 10-III usually
double (1-2); 3-VI double; 4,10,12-VII double. Segment VIII: Comb scales 55 to a-
bout 70; free portion of midapical scale shorter than sessile portion, length 0.017-
0.021 mm (0.019-0.021); hair 2-VIII usually double (2-3). Siphon: Length 0.61-
OFS mm, L/S 2:38-2:86 (213-2 86). P/L 048-052 “(046-0 53). Hil 0 54-0 57
(0.54-0.58); hair 8-S single. Anal Segment: Saddle usually extending less than half-
way around segment; ventral brush usually with 10 hairs (10-11); 4a-X usually 8-
branched (7-9).
SYSTEMATICS. Aedes argyrothorax shows many unique features in the male
genitalia. In addition to the foliaceous claspette filament and laterally declivous
margin of the mesal lobe of the prosophallus, mentioned in the description, there
are the following characters: (1) sidepiece with sternomesal convexity subapical in
position, (2) clasper not markedly tapered distally, the axis distinctly curved, (3)
median lobe of prosophallus virtually obsolete and mesal lobe greatly reduced lat-
erally (consequently declivous laterad), and (4) paraproct with several small teeth
at apex. The male is further distinguished by the very short palpus and the broad
and only slightly emarginate anterior silver marking of the mesonotal disc. The larva
is characterized by the hairs generally with few branches, hair 8-S, in particular, be-
ing single (multiple in the other species); also, the anal saddle is more widely incom-
plete than in the other species.
A series of females from Iquitos, Peru, are treated here as argyrothorax. Their
identity, however, remains in doubt since the male is not represented and since all
other records of this species are from coastal areas.
Cerqueira (1943:32), Martinez (1950:38) and Prosen, Carcavallo and Martinez
(1964:102) have reported argyrothorax from the department of Santa Cruz in Bo-
livia, far south of the confirmed range of the species. These records probably repre-
sent a species of the Terrens Subgroup rather than argyrothorax.
DISTRIBUTION (fig. 2). Coastal lowlands from eastern Venezuela south to state
of Rio de Janeiro, Brazil; Iquitos, Peru, based upon questionable determination. Ma-
terial examined: 78 specimens; 11 6, 22 9, 18 pupae, 27 larvae; 8 individual rearings
(6 larval, 1 pupal, 1 incomplete).
BRAZIL. Amapa: Macapa [elev. near sea level] , May 1948 (1587), 2 Ip? (1587-5 9), 1 $(1587-
1), 1 Ip (458, adult number unknown) [USNM]. Ceara: Serra da Pacoti (Cerqueira 1950:177). Gu-
anabara: Rio de Janeiro, Feb 1938, 1 ¢6 [USNM]. Para: Cameta [elev. near sea level] (Cerqueira
1950:177). Curralinho [elev. near sea level] , Feb 1936, 1 6 (1178) [USNM]. Igarape-Miri [elev. O-
38 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
300 ft] and Nova Timboteua (Cerqueira 1950:177). Pernambuco: Recife [elev. near sea level]
(Cerqueira 1950:177). Rio de Janeiro: Mangaratiba [elev. 0-300 ft] , Dec. 1938, 2 6 [USNM].
FRENCH GUIANA. Guyane: Cabassou [elev. 0-100 ft], 31 Jan 1965, small treehole, height 1-3
ft, T. Aitken, R.Martinez, A. Guerra (FG 12,14), 1 Ipd (12-11), 3 Ip? (12-12,13; 14-12), 1 p?
(14-11), 1 6, 3 p (12-10), 1 6, 4 p, 7 L(12-1), 13 L(14-1) [UCLA] . Inini: Langa Tabiki (Maroni
River, ca 75 km S Saint Laurent) [elev. 0-300 ft], Oct 1947, 1 6 (FGA 46-1) [UCLA] . Political
subdivision not specified: Maroni River, Oct 1947, 1 3, 6 9 (FGA 45-1) [UCLA]. Locality not
specified: 1944, H. Floch, 1 9 [UCLA].
GUYANA. North West: Mabaruma [elev. 0-300 ft] , 22 Dec 1945, Anne Peberdy, 3 ? (207E-
39) [UCLA].
PERU. Loreto: Iquitos [elev. 300-700 ft] , 5 9, R. Shannon [USNM].
SURINAM. Nickerie: Corentyne River (Corantijn; probably vicinity of Oreala or Epira, Guy-
ana) [elev. 0-300 ft], 1 July 1907, J. Aiken, 1 ? (DG-3) [USNM]. Suriname: Paramaribo [elev.
near sea level], in house, 1 6 [USNM 22713]. Geiersvlijt (near Paramaribo) [elev. near sea level]
(holotype) [ITH].
VENEZUELA. Delta Amacuro: Manoa [elev. near sea level], 10 Jan, F.L. de V., 2 29. Manoa
Woods, 10 Jan, F.L. de V., 1 9 [USNM].
Bertrami Subgroup
ADULT (female only). Head: Vertex with decumbent scales along longitudinal
midline narrow; occiput with erect scales pale; proboscis subequal in length to fe-
mur I. Thorax: Mesonotal disc with very broad anterior silver band; complete acrost-
ichal or posterior dorsocentral lines absent; acrostichal setae absent; fossal macula
coextensive with fossa; supraalar macula truncate posteriorly, broadly joined to fos-
sal macula; ppn silver scaled; ssp scales present; pra hairs predominantly pale; upper
stp and mep scale patches not contiguous. Legs: Mid- and hindlegs not shaggy; mid-
and hindtarsi with conspicuous silver markings; tarsus 1-I with narrow apical silver
band; femur II with knee spot broad, the silver scales extending basad of anterior
subapical setae; tarsus 1-I] with median dark band complete, narrow; tarsus 2-II with
complete apical dark band; tarsus 5-II without silver scales; femur III with basal dark
band incomplete; tarsus 5-III without silver scales. Wing: Vein C with small basal
patch of silver scales.
PUPA, LARVA. Unknown.
DISCUSSION. The monobasic Bertrami Subgroup, known only by a single adult
female, is distinguished from the other subgroups by the very broad anterior band
of the mesonotum. The supraalar macula is posteriorly truncate as in the Thorntoni
Subgroup.
The form of the supraalar macula suggests a phyletic relationship of the Bertrami
and Thorntoni Subgroups. The transversely silvered mesonotal disc and the lowland
distribution in British Honduras just to the north of the northern limits of thorn-
toni in the coastal lowlands of Central America (fig. 2) is further suggestive of a
relationship with the Thorntoni Subgroup.
3. Aedes (Finlaya) bertrami Schick, n.sp.
Figs. 2,11
TYPE: Holotype ° (BH 395-2), new capital site, 0.6 mi above jct of Belize-Cayo Road and
Schick: Terrens Group of Aedes (Finlaya) 39
Hummingbird hwy, Cayo, British Honduras [elev. 200 ft] , 8 Aug 1967, biting-landing, 1800-1830
‘hrs, D.S. Bertram [USNM].
FEMALE. Head: Decumbent scales laterad of median longitudinal line broad,
forming dark macula bounded by silver and clear scales. Thorax (fig. 11): Meson-
otal disc with anterior silver band equal in length to fossa; acrostichal line absent;
prescutellar space and scutellum with silver scales. Legs: Femora I and II with well-
developed posterior patch of silver scales in basal half, subequal in size, median dark
band of tarsus 1-II about 0.25; femur III with basal dark band incomplete, very
poorly developed, subapical dark band 0.22; tarsus 1-III with basal silver band 0.09,
apical silver band 0.26. Wing: Vein R without silver scales.
MALE, PUPA, LARVA. Unknown.
DISTRIBUTION (fig. 2). Coastal lowlands of British Honduras. Known only by
holotype female; no individual rearings.
Terrens Subgroup
ADULTS. Head: Vertex of both sexes with decumbent scales along longitudinal
midline narrow; occiput of females with all erect scales pale or with mixture of pale
and dark scales, of males with all erect scales pale; palpal segment 3 of males with
several long apical ventrolateral hairs forming tuft as long as segments 4 and 5 com-
bined; segment 4 with hairs of ventrolateral row long, closely spaced. Thorax: Meso-
notal disc of females not transversely silvered, of males with narrow to moderately
broad anterior silver band or not transversely silvered; complete acrostichal line
rarely present and posterior dorsocentral line absent in females; acrostichal setae
present or absent; fossal macula of females not coextensive with fossa, of males
coextensive or not coextensive with fossa; supraalar macula not truncate posteriorly,
in females broadly joined to fossal macula; ppn silver scaled; ssp scales present or
absent; pra hairs of females pale or dark; upper stp and mep scale patches not con-
tiguous. Legs: Mid- and hindlegs not shaggy; mid- and hindtarsi with conspicuous
silver markings; tarsus 1-I with or without narrow apical silver band; femur II with
knee spot broad, the scales extending basad of anterior subapical setae; tarsus 1-II
with median dark band usually complete and moderately broad; tarsus 2-II with
complete apical dark band; tarsus 5-II without silver scales; femur III with basal
dark band complete, broad; tarsus 5-III without silver scales. Wing: Vein C of fe-
males with small basal patch of silver scales or line reaching crossvein h, of males
with line reaching h; vein R of males with small basal patch of silver scales or line
reaching level of h; vein Cu of males without silver scales.
MALE GENITALIA. Sidepiece: Basal tergomesal area without dense patch of
long setae; median sternomesal area without strongly developed sclerite and with-
out tuft; specialized subapical sternal seta absent. Prosophallus: Mesal lobe with
lateral portion usually not inclined or inclined at 15° or less from horizontal; stem
not bowed; filament with hook not strongly angulate.
PUPAE. Cephalothorax: With or without pale inverted V-shaped marking; hair 5-
C less than twice length of 4-C; 9-C usually single. Abdomen: Hair 1-I with primary
branches predominantly multiple; 2-II usually not laterad of 3-II, usually mesad of
3-II for 0.3 or more the distance from 1-II to 3-II; 3-III usually single. Paddle: Pig-
mented; apex not or at most weakly produced; hair 1-P shorter than paddle.
LARVAE. Head: Hair 5-C usually single, at most triple; 6-C usually single, at most
40 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
double; 7-C more than half length of 6-C; 11-C longer than mental plate; 14-C about
subequal in length to or longer than mental plate, single; bmh single. Antenna: Hair
1-A single. Thorax: Hair 11-P less than half length of 14-P; 14-P usually single; 3-M
usually single; 4-M double to multiple; 8-T shorter than but more than half length of
metathoracic pleural tubercle. Abdomen: Hair 5-I at least subequal in length to 4-
I; 2-II well mesad of 4-II; 3-III double to multiple; 10-III usually single; 3-VI usu-
ally single; 4-VII branched; 5-VII caudad of 4-VII, usually less than half length of,
or at most about equal to half length of 3-VII; 10,12-VII single or branched. Seg-
ment VIII: Midapical comb scale with free portion shorter than or subequal in
length to sessile portion; hair 2-VIII single or branched. Anal Segment: Saddle ex-
tending around segment for moderate distance, ventral margin without deep slit.
DISCUSSION. Five species are recognized in the Terrens Subgroup, terrens, bra-
ziliensis, zavortinki, apollo and berlini. The adults are difficult to characterize be-
cause of the great amount of variation and the few characters that are constant do
not serve to separate the subgroup from some of the others. The adults can be rec-
ognized in general by the combination of the narrow curved scales along the longi-
tudinal midline of the vertex, the anterior silver band on the mesonotal disc of the
male, the broadly joined fossal and supraalar maculae, the complete and broad band
of femur III and the tarsi which show the typical banding pattern of the group.
However, only the characters of the vertex scales, joined maculae and tarsal band-
ing pertain to all the species. The male genitalia and pupae do not show distinctive
subgroup characters. The larvae are characterized by the combination of the single
to triple hairs 5,6-C, the normal length of 7,11-C, 8-T and 5-I (as opposed to the
short condition as in the Thorntoni Subgroup) and the branched 4-VII.
The Terrens Subgroup is the dominant subgroup on the Atlantic side of South
America and is distributed from the Canal Zone southward into Bolivia, northern
Argentina and southeastern Brazil. As suggested in the chapter on Systematics and
Evolution, this subgroup is rather primitive and represents the ancestral stock of
the southern subgroups of the Terrens phyletic line that occur in the Atlantic drain-
ages of Panama and South America.
The treatment of the Terrens Subgroup must be considered as quite preliminary.
The female of braziliensis and male of berlini are unknown and the identity of bra-
ziliensis is in doubt. More material is necessary to properly determine and charact-
erize the species found in northeastern South America because of the complexity
of the subgroup in the region. Many of the specimens cannot be satisfactorily
placed with the known species and may represent variants of these species, hy-
brids or unrecognized species (see terrens and apollo). More material is also nec-
essary to properly characterize terrens in Brazil because of the considerable varia-
tion of this species. Besides the possibility of more species existing in northeastern
South America, 2 or 3 undescribed forms are present in northwestern Argentina
and probably in Bolivia (see argyrothorax and terrens).
Aedes braziliensis, zavortinki, apollo and possibly terrens appear to form a spec-
ies complex, zavortinki and apollo probably being the most closely related. Aedes
berlini stands apart from the others because of the absence of ssp scales and the
long hair 7-VII of the pupa. Perhaps this species should be set aside as a separate
subgroup; the discovery of the male should help clarify its placement.
Schick: Terrens Group of Aedes (Finlaya) 41
4. Aedes (Finlaya) terrens (Walker)
Pigs. 3.7.8 1516
1856. Culex terrens Walker, 1856:429. TYPE: Holotype 6 with attached genitalia mount, South
America [BM]. Type data from Belkin (1968:8).
1904. Gualteria oswaldi Lutz In Bourroul, 1904a:13;1904b:4; Lutz, 1905:65-66. TYPE: Adults,
states of Rio de Janeiro (? in part Guanabara) and Sao Paulo, Brazil, “sylvan, principally
in the forests of the mountains” [translation mine] [BM, other material possibly in IOC
or FMP]. Type data from Belkin (1968:6). Synonymy with terrens by Dyar 1921:152.
Aedes (Finlaya) terrens of Shannon (1931a:8,23); Duret and Barreto (1956:90); Belkin (1968:8).
Aedes terrens of Bonne-Wepster and Bonne (1921:23); Davis and Shannon (1931:716); Shannon,
Whitman and Franca (1938:110-111); Antunes and Lane (1938:1036); Causey, Kumm and
Laemmert (1950:302,304); Alvarado, Coll, et al (1959:194); Causey, Causey, et al (1961:235).
Aedes (Finlaya) terrens in part of Dyar (1921:152-153; 1928:224-225); Bonne-Wepster and Bonne
(1925 :424-428); Shannon (1931b:147-148); Edwards (1932:150); Lane (1939:104-105; 1953:
686-687); Horsfall (1955:462); Stone, Knight and Starcke (1959:171); Forattini (1965 :395-
396).
Aedes (Gualteria) terrens of Vargas (1950a:62).
Stegomyia terrens of Theobald (1901:305-306; 1910:174); Giles (1902:378); Lutz (1905:66);
Surcouf and Gonzales-Rincones (1911:135).
Culex terrens of Giles (1900:241); Theobald (1901:423; 1905:26).
Aedes (Finlaya) oswaldi of Belkin (1968:6).
Aedes oswaldi of Howard, Dyar and Knab (1917:815-819); Dyar (1918:73,80); Bonne-Wepster
and Bonne (1920:23-24).
Haemagogus oswaldi of Dyar and Knab (1906a: 166).
Gualteria oswaldi of Blanchard (1905:633); Theobald (1907:552-554); Peryassu (1908 :45 ,64,177-
179); Theobald (1910:606); Surcouf and Gonzales-Rincones (1911:149).
Aedes (Finlaya) terrens var. podographicus of Costa Lima (1930:257-258); Lane (1936a:11).
Aedes terrens var. podographicus of Lane (1936b:131).
FEMALE (figs. 7,15). Head: Vertex with decumbent scales laterad of median
longitudinal line dark, narrow curved; occiput with erect scales usually all pale.
Thorax (fig. 15): Acrostichal line usually absent (complete line present in Teresopo-
lis, Brazil); acrostichal setae usually absent, when present most posterior seta usu-
ally at about 0.25 from anterior end, rarely as far caudad as 0.5; fossal macula
separated from lateral margin of fossa by a variably developed but usually prom-
inent area of dark scales, infrequently by only a few dark scales, often divided
into anterior and posterior portions; prescutellar space and scutellum with or with-
out silver scales; ssp scales present; pra hairs usually pale (dark in 1 specimen
from Londrina, Brazil). Legs: Femora I and II with well-developed posterior patch
of silver scales in basal half, that of femur II often larger; tarsus 1-I] with med-
ian dark band complete, about 0.33-0.5; femur III with basal dark band 0.11-0.17,
subapical dark band 0.24-0.33 (0.17-0.33); tarsus 1-III with basal silver band 0.07-
0.09 (0.04-0.10), apical silver band 0.19-0.25 (0.16-0.26). Wing: Vein C with silver
scalation at base from small patch to line reaching crossvein h; vein R without silver
scales or with a small basal patch. '
MALE (figs. 8,15). Head: Vertex with decumbent scales laterad of median longi-
tudinal line silver, generally narrow curved; palpus subequal in length to or 1 lab-
ellum length shorter than proboscis. Thorax (fig. 15): Mesonotal disc usually with-
out anterior silver band, rarely with very narrow band a single row of scales in
42 Contrib. Amer. Ent. Inst. vol. 5, no. 3, 1970
width (Rio Cururipe, Brazil) and with or without strip of silver scales bordering
fossal macula; fossal macula usually not reaching mesal margin of fossa, but in Salva-
dor region of Brazil often coextensive with fossa (in latter case strip of silver scales
bordering macula present); pra hairs as in female. Legs: Tarsus 1-I] with median
dark band variably developed [in Salvador region and occasionally farther south in-
complete or narrow, less than 0.33; in southern Brazil usually complete, at least
about 0.33]. Wing: Vein R with silver scalation at base forming small patch or
interrupted line reaching level of crossvein h.
MALE GENITALIA (fig. 15). Sidepiece: Length 0.31-0.38 mm. Prosophallus:
Length 0.09-0.13 mm; width 0.11-0.14 mm; mesal lobe with lateral portion in Sal-
vador region of Brazil usually inclined less than 15° from horizontal, sometimes
not appreciably inclined, and in only genitalia from other locality, Cambara, Brazil,
between 15° and 30°; stems usually divergent, sometimes parallel, filament ratio
0.51-0.84 (0.42-1.07; 2 highest values, 1.00 and 1.07, possibly due to abnormal in-
clination of filament). Aedeagus: Length 0.13-0.14 mm.
PUPA (fig. 15). Cephalothorax: Without pale inverted V-shaped marking. Abdo-
men: Hair 2-II mesad of 3-II for 0.4 or more the distance from 1-II to 3-II; 3-II]
single; 7-VII not reaching caudolateral margin of sternite VIII. Paddle: Not tapered;
apex not produced; ventral midrib weakly pigmented for most of the length, more
strongly at apex. |
LARVA (fig. 16). Head: Hair 5-C usually double or triple (1-4); 6-C usually
single or double, infrequently triple; 7-C triple to 9-branched [in Novo Iguassu,
Brazil, triple to 5-branched; in other localities 6-9 branched]; 14-C longer than
mental plate, particularly long in Anapolis, Brazil. Thorax: Hair 1-P usually double
or triple (2-4); 4-P usually double or triple (1-4); 5-P usually double (2-3); 7-P usu-
ally triple (2-4); 14-P single; 3-M usually single (1-2); 4-M double or triple. Abdo-
men: Hair 3-III double to 4-branched; 10-III single, infrequently double; 3-VI sing-
le or double; 4-VII double to 4-branched; 5-VII usually less than half length of 3-
VII, somewhat longer than half length in Cha Pilar, Brazil; 10-VII usually double
(1-2); 12-VII single to triple. Segment VIII: Comb scales 33-85; length of free por-
tion of midapical scale 0.022-0.026 mm; hair 2-VIII usually double (1-2). Siphon:
Length 0.75-1.02 mm; L/S 1.96-2.28; P/L 0.44-0.45: H/L 0.48-0.62. Anal Segment:
Ventral brush usually with 14 hairs (13-15); 4a-X usually 8-branched (6-11).
SYSTEMATICS. Brazilian specimens of terrens show some marked variation in
characters that are generally of a specific nature in the Terrens Group, in the fe-
male, the presence or absence of a complete acrostichal line and the development
of either pale or dark pra hairs, and in the male the fossal macula coextensive or
not coextensive with the fossa. The mep scale patch may be entire or transversely
divided, the latter state being unique for the group.
There are populations of the Terrens Subgroup from northeastern South America
whose identities are uncertain but which show some similarities to terrens. Among
some females from the Maracay region of Venezuela, 2 are strikingly similar to those
Brazilian specimens of terrens which show the characteristic sublateral fossal stripe
(fig. 7); the pra hairs, however, are dark unlike those of typical terrens. The other
females, perhaps conspecific with these 2, show a macula that is broadly developed
up to the lateral margin of the fossa, a condition I have not encountered in Brazilian
terrens. | hesitate to treat the 2 females as terrens since, among other possibilities,
they may represent variants of a form found on the coast of Venezuela which is
apparently related to apollo (see). More than 1 species may be represented in several
collections from French Guiana (all probably from the Ile de Cayenne). The females
Schick: Terrens Group of Aedes (Finlaya) 43
are similar to terrens except that the fossal macula is broadly developed up the lat-
eral margin of the fossa. The mesonotum of the males is similar to that of the terrens
from Rio Cururipe, Brazil (fig. 15), in that a narrow anterior mesonotal band is pres-
ent and the fossal macula is coextensive with the fossa, but in some of the males
the band is broader, about 0.25. The larva of 1 of these forms shows a triple hair 4-
VII and double 2-VIII as in terrens. In northwestern Argentina terrens is apparently
largely replaced by a form in which the fossal macula is absent, the basal band of
femur III is incomplete and in which there are no silver scales at the base of wing
vein C of the female. Specimens of this form came to my attention too late to be
formally treated in the present revision.
DISTRIBUTION (fig. 3). Brazil, south of Amazon basin, Paraguay and north-
ern Argentina, predominantly at elevations of 700-3300 ft. Material examined: 116
specimens; 29 6, 43 9, 13 pupae, 31 larvae; 12 individual rearings (4 larval, 4 pupal,
4 incomplete).
ARGENTINA. Misiones: Iguazu [elev. 0-700 ft], 18 June 1927, R. Shannon, E. Del Ponte, 1 ?
(217) [USNM] . Tucuman: Tafi Viejo [elev. 1600-3300 ft] , 14 Mar 1927, R. Shannon, E. Del Pon-
te, 1 9 [USNM].
BRAZIL. Alagoas: Cha Pilar [elev. 700-1600 ft] , 1 L (2398) [USNM] . Bahia: Cururipe (? Cor-
uripe in Alagoas), 28 Sept 1920, 1 6, 1 ? [USNM]. Piraja, 27 Dec 1928, treehole, N. Davis, R.
Shannon, 1 p (27XII1), 1 1 (27XID): 7 Feb 1929, treehole, N. Davis, R. Shannon, 2 Ipd (14112,
1SI1), 1 Ip? (15113), 2 (13111, 14113), 2 Ip (13111, 15112), 1 1(14111), 16, 1 9: 1 Mar 1929, tree.
hole, N. Davis, R. Shannon, 1 Ip (7IID), 1 1 (212); 13 Mar 1929, bamboo, N. Davis, R. Shannon, 1
Ip (28-3-1): 10 Apr 1929, Bahioe. R. Shannon, | 6; 11 Apr 1929, treehole. R. Shanon: i's, 2 3
genitalia (28V6, 13); 16 July 1929, treehole, R. Shannon, 1 1° (24VII2), 10 1 (19VII1,2; 22VII1-6;
23VII1; 24VII1); Jan 1930, R. Shannon, 2 é: 1930; 1:2 [USNM]. Rio Cururipe (? Rio Coruripe ; in
Alagoas), 13 Jan 1931, H. Kumm, 1 3 (BM 1933- 503) [BM]. Locality not specified: 1930,1 6,1 9
[USNM]; H. Kumm, f 3 (BM 1933 -503) [BM]. Goias: Anapolis [elev. 3300-6600 ft] , Mar 1936,
1 2 (6989); date not specified, 5 5 (6600,7131,6844,6989,15389), 7 L (5557) [USNM] . Guana-
bara: Parque de Gavea (767), 1 6 (767-37), 2 9, 2 p, 21(767-56,57) [USNM] . Rio de Janeiro, 21
Jan 1907, 1 2 (1034), Dec 1937, 4 9 [USNM] . Mato Grosso: Maracaju [elev. 1600-3300 ft] , July
1937, 1 2; Nov 1937, 10 2; Dec 1937, 6 9; June 1938, 1 9; date not specified, 1 6, 2 9 [USNM].
Minas Gerais: Araxa [elev. ca 3300 ft], 2 L (26299) [USNM]. Parana: Cambara [elev. 700-3300
ft], Sept 1936, 4 5 (11493); date not specified, 1 6 (11487) [USNM]. Londrina [elev. 700-1600
ft], Feb 1937, 1 9 (12458) [USNM]. Rio de Janeiro: Nova Iguacu [elev. 0-300 ft], 3 L (3174)
[USNM]. Teresopolis [elev. 1600-3300 ft], Nov 1942, 1 9(12/28774); Dec 1942, 1 6 (1/29078)
[UCLA]. Sao Paulo: Cantareira..., Jan 1944, treehole (1277), 1 pd (1277-31), 3 pe (1277-8 ,12,
15) [USNM]. Rio Buri (533) [USNM]. Rocinha, 20 Feb 1937, human bait in forest, A. Ramo, 1
9 (744) [USNM].
PARAGUAY. Cordillera: San Bernardino [elev. 700-1600 ft] , Fiebrig, 1 9 [USNM].
5. Aedes (Finlaya) braziliensis Gordon & Evans
Figs. 3,21
1922. Aedes (Finlaya) oswaldi var. braziliensis Gordon and Evans, 1922:329. TYPE: Holotype
3 (10.1/463), Macapa (about 15 mi from Manaus), Amazonas, Brazil, 8 Dec 1921, rear-
ed from larva ex treehole, R.M. Gordon [BM]. Type data from Belkin (1968:4).
Aedes (Finlaya) braziliensis of Belkin (1968:4).
Aedes (Finlaya) terrens var. braziliensis of Bonne and Bonne-Wepster (1925:427).
Aedes (Finlaya) terrens in part of Dyar (1928:224); Edwards (1932:150); Lane (1939:104-105;
1953:686-687); Knight and Marks (1952:549); Stone, Knight and Starcke (1959:171); Forat-
tini (1965:395).
Ad. Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
FEMALE. Unknown.
MALE. Head: Vertex with decumbent scales laterad of median longitudinal line
dark, broad; palpus less than 1 labellum length shorter than proboscis. Thorax (fig.
21): Mesonotal disc with anterior silver band about 0.75 length of fossa, with strip
of silver scales bordering fossal macula; acrostichal setae absent; fossal macula co-
extensive with fossa; ssp scales present; pra hairs dark, more heavily pigmented than
those of ppl or upper mep. Legs: Femur I without posterior patch of silver scales;
femur II with posterior patch comprising only a few scattered scales; tarsus 1-II with
median dark band complete, about 0.33; femur III with basal dark band 0.20, sub-
apical dark band 0.26; tarsus 1-III with basal silver band 0.05, apical silver band
0.23. Wing: Vein R with basal line of silver scales reaching about 0.5 to level of h.
MALE GENITALIA (fig. 21). Sidepiece: Length 0.34 mm. Prosophallus: Length
0.10 mm; width 0.13 mm; mesal lobe with lateral portion inclined at about 15 oor
somewhat less, from horizontal; stems divergent; filament ratio 1.05 (this high value
possibly due to abnormal inclination of filament). Aedeagus: Length 0.12 mm.
PUPA, LARVA. Unknown.
SYSTEMATICS. As no topotypic material of braziliensis was available for this
study the description of the braziliensis here is based upon one male specimen,
“French Guiana” (671016-12). A second male (FG 14-13), from Cabassou, Ile de
Cayenne, was in poor condition and not used in the description. There is some
doubt about the identity of this material with topotypic braziliensis. In his study
of the types of New World species in European museums, John N. Belkin exam-
ined the holotype male of braziliensis, the genitalia of which was missing. His un-
published sketch of the mesonotum matches that of the French Guianan speci-
mens and is the basis of the present identification. Belkin’s notes also indicate agree-
ment in other minor respects. Discrepancies lie in the widths of the basal silver band
of tarsi 2-II and 2-III, about 0.5 and 0.25 respectively in the type of braziliensis and
about 0.33 for both segments in the French Guianan specimens. These discrepan-
cies are probably not significant since the extent of the markings is subject to a sim-
ilar range of variation in other species. The significance of the “capitate” clasper
and the dorsally dark-scaled abdominal segment VIII that Gordon and Evans cite
as separating braziliensis from oswaldi (terrens) is unknown.
Aedes zavortinki and apollo have mesonotal markings similar to those of the
French Guianan specimens but it is less likely that these would be braziliensis since
their ranges, admittedly very incompletely known, lie in or beyond the highlands
of Colombia. The type locality of braziliensis is from near Manaus, Brazil.
Aedes braziliensis, as treated here, differs from the other species of the subgroup
by a combination of the broad lateral decumbent scales of the vertex and the dark
pra and upper mep hairs.
DISTRIBUTION (fig. 3). French Guiana and the state of Amazonas, Brazil. Mat-
erial examined: 2 6; no individual rearings.
BRAZIL. Amazonas: Macapa (nearest town Manaus) [elev. 0-300 ft] , 8 Dec 1921, treehole, R.
Gordon (type series).
FRENCH GUIANA. Guyane: Cabassou, elev. ca 100 ft, 31 Jan 1965, small treehole, height
2 ft, T. Aitken, R. Martinez, A. Guerra, 1 6 (FG 14-13) [UCLA]. Locality not specified: 1944, H.
Floch, 1 6 [UCLA].
Schick: Terrens Group of Aedes (Finlaya) 45
6. Aedes (Finlaya) zavortinki Schick, n.sp.
Figs. 3,17,18
TYPE: Holotype 6 (207C-38) with associated larval skin (5-163), Barro Colorado Island [elev.
100-500 ft] , Canal Zone, Panama, 22 May 1945, large treehole, WH.W. Komp [USNM] . Allotype
? (207D-32) with associated larval and pupal skins (5-181), same data as holotype [USNM]. Para-
types: 1 6, 2 2,3 p, 2 1(207D-43, 5-182), 7 6 (207C-10), 3 6 (207C-8,14,42), 2 6 (207C-45), 1
d (207C-34), same data as holotype [BM; USNM; UCLA]. This species is dedicated to Thomas J.
Zavortink.
Aedes thorntoni in part of Howard, Dyar and Knab (1917:819-821); Dyar (1918:225-226).
Aedes insolitus in part of Busck (1908:64).
FEMALE. Head: Vertex with decumbent scales laterad of median longitudinal
line generally dark, narrow curved; occiput with erect scales pale. Thorax (fig. 17):
Acrostichal setae usually absent, when present most posterior seta at about 0.25
from anterior end; fossal macula fully developed up to lateral margin of fossa:
prescutellar space and scutellum with or without silver scales; ssp scales present;
pra hairs dark, more heavily pigmented than those of ppl or upper mep. Legs: Fe-
mora I and II with well-developed posterior patch of silver scales in basal half, that
of femur II usually somewhat larger; tarsus 1-II with median dark band complete,
about 0.33-0.5; femur III with basal dark band 0.13-0.21, subapical dark band 0.26-
0.33; tarsus 1-III with basal silver band 0.11-0.12 (0.10-0.17), apical silver band
0.20-0.28 (0.19-0.31). Wing: Vein C with basal silver scales forming small patch
or line reaching crossvein h; vein R without silver scales or with small basal patch.
MALE. Head: Vertex with decumbent scales silver, narrow curved; palpus about
1-2 labellum lengths shorter than proboscis. Thorax (fig. 17): Mesonotal disc with
anterior silver band about 0.25-0.5 length of fossa and with strip of silver scales
bordering fossal macula; fossal macula coextensive with fossa: pra hairs as in fe-
male. Legs: Tarsus 1-II with median dark band complete, at least 0.33. Wing: Vein
R with basal silver scalation forming small patch or interrupted line reaching level
of crossvein h.
MALE GENITALIA (fig. 17). Sidepiece: Length 0.32-0.35 mm (0.27-0.42 mm).
Prosophallus: Length 0.11-0.12 mm (0.10-0.14 mm); width 0.14-0.15 mm (0.13-
0.15 mm); mesal lobe with lateral portion usually inclined at about 15° or less
from horizontal (sometimes between 15° and 30°, infrequently not appreciably
inclined or as great as 30°); stems usually convergent, sometimes parallel, infrequent-
ly divergent; filament ratio 0.51-0.66 (0.42-0.77). Aedeagus: Length 0.13-0.14 mm.
PUPA (fig. 17). Cephalothorax: Without pale inverted V-shaped marking. Abdo-
men: Hair 2-II rarely laterad of 3-II, usually mesad for 0.3 or more the distance
from 1-II to 3-II; 3-III usually single, rarely branched; 7-VII not reaching caudo-
lateral margin of sternite VIII. Paddle: Not tapered; apex not produced: ventral
midrib weakly to moderately pigmented for most length, not more strongly at apex.
LARVA (fig. 18). Head: Hair 5-C usually double (1-3); 6-C single; 7-C usually 6-7
branched (5-9); 14-C slightly shorter than or subequal in length to mental plate.
Thorax: Hairs 1,4,5-P double or triple; 7-P usually triple (2-4); 14-P single; 3-M sing-
le; 4-M triple. Abdomen: Hair 3-III triple; 10-III single; 3-VI single; 4-VII double; 5-
VII less than half length of 3-VII; 10-VII single or double; 12-VII usually double, in-
frequently single. Segment VIII: Comb scales about 75; length of free portion of
midapical scale 0.026-0.035 mm; hair 2-VIII single. Siphon: Length 0.80-0.97 mm;
46 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
L/S 2.08-2.44; P/L 0.45-0.50; H/L 0.51-0.57. Anal Segment: Ventral brush usually
with 15 or 16 hairs (13-16); 4a-X usually 10-12 branched (9-12).
SYSTEMATICS. Aedes zavortinki is distinguished from the other members of
the subgroup in the different stages only by combinations of characters, in the fe-
male by the narrow curved lateral decumbent scales of the vertex, the relatively
broad fossal macula, the presence of ssp scales and the dark pra but pale upper
mep hairs; in the male by the same vertex scale and pleural hair characters and by
the anteriorly silvered mesonotal disc; and in the larva by the usually double hair
5-C, at least 5-branched 7-C, single 2-VIII, ventral brush of 15 or 16 hairs (usually
the latter) and the 10-12 branched 4a-X.
DISTRIBUTION (fig. 3). Canal Zone and Pacific lowlands of eastern Panama.
Material examined: 158 specimens; 57 6, 52 9, 22 pupae, 27 larvae; 16 individual
rearings (15 larval, 1 pupal). For the Komp collections see the explanatory chapter.
PANAMA. Canal Zone: Barro Colorado Island [elev. 100-500 ft], 15 May 1945, treehole, W.
Komp, 2 6 (207C-3), 1 1 (207D-30) — 1 6, 1 9 (207C-11), 1 6 (207C-24) [UCLA; USNM]; 22
May 1945, large treehole, W. Komp, type series, 1 16 (207C-38), 1 lp? (207D-32), 1 6,2 9,3 p,2
1 (207D-43), 7 6 (207C-10), 3 b (207C-8,14,42), 2 3 (207C-45), 1 6 (207C-34) [USNM; BM;
UCLA] — treehole, 1 1d (207E-20), 2 lp? (207D-36; 207E-28) — 2 6 (207C-39) — treehole, 1 9, 1
p, 3 1(207D-28) — treehole, 1 lp? (207D-40) — treehole, 2 ¢, 2 1(207E-13) — treehole, 1 ¢ (207E-
26) — deep treehole, 1 Ip? (207E-29); 23 May 1945, treehole, W. Komp, 5 lp? (207D-16; 207E-
18,21,22,27), 1 19 (207D-31), 2 9, 2 p, 21(207E-24) — treehole, 1 Ip? (207D-27); 25 June 1945,
W. Komp, | 6, 2 2,3 p, 3 1(207D-45) [UCLA; USNM]. Camacho, 14 Jan 1922, J. Shropshire, 2 9;
17 Mar 1922, J. Shropshire, 2 6, 3 9; 22 Apr 1922, J. Shropshire, 2 d, 2 9; 1 June 1922,
J. Shropshire, 10 6, 4 9 [USNM]. Chiva Chiva, elev. 0-300 ft, 11 Nov 1965, treehole, height 4 ft,
A. Quinonez, 2 6, 2 p(PA 773-3) [UCLA]. Corozal [elev. 0-100 ft] , 13 Jan 1943, treehole, 1 6, 2
2 (207B-22) [UCLA]. Corozal Hospital grounds [elev. 0-100 ft], 7 Sept 1943, treehole, H.
Herman, | 1° (207B-8) [UCLA; USNM]. Empire [elev. 200-300 ft] , Jan 1922, J. Shropshire, 1 9;
4 Apr 1922, J. Shropshire, 4 6; 10 May 1922, 3 6, 2 9; 21 Aug 1923, H. Dyar, R. Shannon,
1 2 (P56) [USNM]. Gamboa, S. trail, 9 June 1943, Elton, 2 d (207B-2) [UCLA]. Gatun [elev.
near sea level], 25 July 1928, D. Curry, 1 9 [USNM]. Largo Remo [elev. near sea level] , 27 July
1926, D. Curry, 1 6; 11 Aug 1926, D. Curry, 1 6, 1 9(15) [USNM] . Mandingo (? Mandinga), 13
Dec 1921, J. Shropshire, 1 6, 3 9; 14 Dec 1921, J. Shropshire, 1 6, 1 9 [USNM]. Paraiso [elev.
0-100 ft], 12 May 1918, treehole, J. Zetek, 1 ¢ (1638), 2 9 [USNM]. Rio Chagres, upper, tree-
hole, A. Busck, 1 6 (102) [USNM]. Summit [elev. 200-300 ft] , 13 Dec 1939, treehole, W. Komp,
1 6 (207E-4); 17 Aug 1941, treehole high in Mango tree, 1 2 (207B-13) [UCLA]. Locality not
specified, 31 Dec 1921, J. Shropshire, 1 @ [USNM] . Darien: Morti, elev. 200 ft, 3 Dec 1966, large
treehole, height 6-9 ft, O. Berlin, R. Hinds, 1 pd (PA 978-100) [UCLA] . Panama: Nuevo Empera-
dor, near, on RT 7, elev. 100-300 ft, 23 Nov 1965, small treehole, height 4 ft, A. Quinonez, 1 ?
(PA 825-2) [UCLA].
7. Aedes (Finlaya) apollo Schick, n.sp.
Figs. 3,19,20
TYPE: Holotype ? (COB 47-20) with associated pupal and larval skins, Finca Vanguardia (on
Villavicencio-Restrepo road, closer to Villavicencio), elev. 1200 ft, 23 June 1965, small treehole,
height 6 ft, E. Osorno-Mesa, Morales, et al [USNM] . Allotype 6, Bosque Ocoa (near Villavicencio,
elev. 1500 ft), 11 June 1964 [USNM]. Paratype: 1 lp? (47-21), same data as holotype [UCLA].
This species is named to commemorate the landing of Apollo 11 on the moon.
? Aedes terrens of Komp (1936:62); Antunes (1937:76).
? Aedes podographicus of Bugher, Boshell-Manrique, et al (1944:31).
Schick: Terrens Group of Aedes (Finlaya) 47
FEMALE. Head: Vertex with decumbent scales laterad of median longitudinal
line generally dark and narrow curved; occiput with mixture of pale and dark
erect scales. Thorax (fig. 19): Acrostichal setae present or absent, when present
most posterior seta at about 0.33 from anterior end; fossal macula fully devel-
oped up to lateral margin of fossa; prescutellar space with or without silver scales;
scutellum without silver scales; ssp scales present; pra and upper mep with hairs dark
and about equally heavily pigmented, more heavily so than those of ppl. Legs: Fe-
mur I with well-developed posterior patch of silver scales in basal half; femur II with-
out posterior patch; tarsus 1-II with median dark band complete, about 0.4-0.5; fe-
mur III with basal dark band 0.21, subapical dark band 0.19-0.25; tarsus 1-III with
basal silver band 0.09-0.11, apical silver band 0.15-0.17. Wing: Vein C with basal
line of silver scales reaching about 0.5 to crossvein h or to crossvein itself; vein R
without silver scales or with only a few at base.
MALE. Head: Vertex with decumbent scales laterad of median longitudinal line
silver, broad; palpus about 0.5 labellum length shorter than proboscis. Thorax: Mes-
onotal disc with anterior silver band about 0.5-0.75 length of fossa and with strip
of silver scales bordering fossal macula; fossal macula coextensive with fossa; pra
hairs as in female. Legs: Tarsus 1-II with median dark band complete, width un-
known. Wing: Vein R with basal silver scalation very variable, when present, form-
ing small patch to line nearly reaching level of crossvein h.
MALE GENITALIA (fig. 19). Sidepiece: Length 0.31-0.34 mm. Prosophallus:
Length 0.11 mm; width 0.11-0.12 mm; mesal lobe with lateral portion not appreci-
ably inclined or inclined at less than 15° from horizontal; stems divergent (based up-
on | specimen); filament ratio 0.47-0.60. Aedeagus: Length 0.12 mm.
PUPA (fig. 19). Cephalothorax: Without pale inverted V-shaped marking. Abdo-
men: Hair 2-II mesad of 3-II for 0.3 or more the distance from 1-II to 3-II; 3-III sing-
le; 7-VII not reaching caudolateral margin of sternite VIII. Paddle: Not tapered; apex
not produced; ventral midrib weakly to moderately pigmented for most length, not
more strongly at apex. |
LARVA (fig. 20). Head: Hairs 5,6-C single; 7-C triple; 14-C slightly longer than
mental plate. Thorax: Hair 1-P double to 4-branched; 4-P single to triple; 5-P single
or double; 7-P double or triple; 14-P single; 3-M single or double; 4-M triple or 4-
branched. Abdomen: Hair 3-III triple or 4-branched; 10-III single; 3-VI single, 4-VII
double; 5-VII less than half length of 3-VII; 10,12-VII double. Segment VIIT: Numb-
er of comb scales not determined, apparently similar to other species of subgroup;
length of free portion of midapical comb scale 0.033 mm; hair 2-VIII single. Siphon:
Length 0.86-0.92 mm; L/S 2.17-2.50; P/L 0.48-0.51; H/L 0.55-0.56. Anal Segment:
Ventral brush with 15 or 16 hairs; 4a-X 9,10-branched.
SYSTEMATICS. Aedes apollo is very similar to zavortinki and may be distin-
guished from that species by the dark hairs of the mep in the adults and by the
single hairs 5,6-C and triple 7-C in the larva.
Two undetermined males from Restrepo and Villavicencio, Colombia, also show
dark hairs on the mep but differ from apollo in that the decumbent scales of the
vertex laterad of the median longitudinal line are narrow curved and the anterior
silver band of the mesonotal disc is narrower (about 0.25). A collection of a similar
form from the coast of Venezuela near Ocumare de la Costa has just come to my
attention. The males agree with the 2 undetermined males from Colombia except
that the anterior silver band of the disc is even narrower (similar to the illustration
of the male terrens from Rio Cururipe, fig. 15). Two of the 3 females show dark
mep hairs, but those of the third female are pale; the fossal macula in all 3, un-
48 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
like that of apollo, is separated from the lateral margin of the fossa by a more or
less well developed area of dark scales (see females from Maracay area of Venezuela ©
mentioned under ferrens). The pupa shows a distinct pale inverted V-shaped mark-
ing on the cephalothorax, unlike apollo, but the larva is quite similar to that of
apollo. Whether these specimens represent variants of apollo or 1 or more distinct
species cannot be determined until reared material from Colombia becomes available.
DISTRIBUTION (fig. 3). Department of Meta, Colombia, at elevations of 1200 to
1600 ft. Material examined: 11 specimens; 3 6, 4 ?, 2 pupae, 2 larvae; 2 individual
rearings (larval).
COLOMBIA. Meta: Bosque Ocoa (near Villavicencio, elev. 1500 ft), 11 June 1944, 1 d, type
series in part [USNM]. Finca Vanguardia (on Villavicencio-Restrepo road, closer to Villavicencio),
elev. 1200 ft, 23 June 1965, small treehole, height 6 ft, E. Osorno-Mesa, Morales, et al, type series
in part, 2 lp? (COB 47-20,21) [USNM; UCLA]. Villavicencio [elev. 1500 ft], 17 May 1939,1 6
(3967); June 1942, W. Komp, 1 2 (207B-45) [USNM]; Jan 1942, W. Komp (H-9-10), 1 3, 1 9
[UCLA]. Villavicencio, river road to Bosque Ocoa [elev. 1500 ft], 1 June 1942, treehole, 1 6
(207B-10) [USNM].
8. Aedes (Finlaya) berlini Schick, n.sp.
Figs. 3,21,22
TYPE: Holotype ? (TOB 131-12), with associated pupal and larval skins, Parrott Hall, 15.5 mile
post, elev. 300-400 ft, nearest town Parlatuvier, Tobago Island, 29 Nov 1965, treehole, R. Martinez
and A. Guerra [USNM]. Paratypes: 2 L (131-1), same data as holotype. Orange Hill, Tobago Is-
land, elev. 200-300 ft, 27 Nov 1965, biting-landing, 1500 hrs, R. Martinez, A. Guerra, 1 9 (TOB
125-1) [UCLA]. This species is dedicated to O.G.W. Berlin.
FEMALE. Head: Vertex with decumbent scales laterad of median longitudinal
line dark, broad; occiput with mixture of pale and dark scales. Thorax (fig. 21):
Acrostichal setae present or absent. most posterior seta at about 0.25 from an-
terior end; fossal macula fully developed up to lateral margin of fossa; prescu-
tellar space and scutellum without silver scales; ssp scales absent; pra hairs dark,
more heavily pigmented than those of pp! or upper mep. Legs: Femora I and II
with well-developed posterior patch of silver scales in basal half, that of femur II
larger; tarsus 1-II with median dark band complete, about 0.33-0.5; femur III with
basal dark band 0.15, subapical dark band 0.18-0.21; tarsus 1-III with basal silver
band 0.08-0.11, apical silver band 0.20-0.26. Wing: Vein C with basal line of silver
scales reaching about 0.5, or somewhat less, to crossvein h; vein R without silver
scales.
MALE. Unknown.
PUPA (fig. 21). Cephalothorax: With poorly defined pale inverted V-shaped mark-
ing. Abdomen: Hair 2-II mesad of 3-II for 0.4 the distance from 1-II to 3-II; 3-Ill
single; 7-VII long, reaching caudolateral margin of sternite VIII. Paddle: Tapered:
apex weakly produced; ventral midrib moderately and uniformly pigmented in distal
third.
LARVA (fig. 22). Head: Hair 5-C double; 6-C single or double; 7-C triple or 4-
branched; 14-C longer than mental plate. Thorax: Hair 1-P triple; 4-P double or trip-
le; 5-P double; 7-P double or triple; 14-P single or double; 3-M single; 4-M double or
triple. Abdomen: Hair 3-II] usually triple (2-3); 10-III usually single (1-2); 3-VI usu-
ally single (1-2); 4-VII double; 5-VII less than half length of 3-VIJ; 10,12-VII single.
Schick: Terrens Group of Aedes (Finlaya) 49
Segment VIII: Comb scales 50-60; length of free portion of midapical scale 0.034
mm; hair 2-VIII usually single (1-2). Siphon: Length 0.84-0.94 mm; L/S 2.44-2.54;
P/L 0.54-0.63; H/L 0.61-0.67. Anal Segment: Ventral brush with 11 or 12 hairs; 4a-
X 6-branched.
SYSTEMATICS. Aedes berlini can be distinguished from the other species of the
subgroup in the female by the absence of ssp scales (the male is unknown but un-
doubtedly also lacks these scales); in the pupa by the very long hair 7-VII (longer
than in all the other species of the Terrens Group); and in the larva by the single
hairs 10,12-VII vs. usually double in the other species, the smaller number of ventral
brush hairs, 11 or 12 (vs. 13 or more), the usually smaller number of branches of 4a-
X, usually 8-12, but as few as 6 (vs. 6), and the usually greater P/L and H/L ratios,
0.54-0.63 vs. 0.44-0.51 and 0.61-0.67 vs. 0.48-0.62.
DISTRIBUTION (fig. 3). Island of Tobago. Material examined: 6 specimens; 2 9,
1 pupa, 3 larvae; 1 individual rearing (larval).
TOBAGO. Orange Hill, elev. 200-300 ft, 27 Nov 1965, biting-landing, 1500 hrs, R. Martinez,
A. Guerra, 1 2 (TOB 125-1, type series in part) [UCLA]. Parrott Hall, elev. 300-400 ft, 29 Nov
1965, large treehole, height 3 ft, R. Martinez, A. Guerra (TOB 131, type series in part), 1 Ip?
(131-12), 2 L (131-1) [USNM; UCLA].
Alboapicus Subgroup
ADULTS. Head: Vertex of both sexes with decumbent scales along longitudinal
midline narrow; occiput of female with erect scales predominantly dark, of male
with all erect scales pale; proboscis of female longer than femur I; palpal segment 3
of male with | or 2 usually short apical ventrolateral hairs; segment 4 with hairs of
ventrolateral row long but sparse. Thorax: Mesonotal disc of female not transversely
silvered, of male with anterior silver band; complete acrostichal or posterior dorso-
central lines absent in female; acrostichal setae absent; fossal macula of female
not coextensive with fossa, reduced mesally, of male coextensive with fossa; supra-
alar macula not truncate posteriorly, in female disjunct from or broadly joined to
fossal macula; ppn silver scaled; ssp scales absent: pra hairs of female dark; upper stp
and mep scale patches not contiguous. Legs: Mid- and hindlegs not shaggy; mid- and
hindtarsi with conspicuous silver markings; tarsus 1-I with or without narrow apical
silver band; femur II with knee spot present, moderately broad, the scales just reach-
ing anterior subapical setae; tarsus 1-II with median dark band complete, moderately
to very broad; tarsus 2-II with complete apical dark band; tarsus 5-II at least partly
silver scaled; femur III with basal dark band complete, usually broad; tarsus 5-III sil-
ver scaled. Wing: Vein C of female with small basal patch of silver scales or line
reaching crossvein h, of male with line reaching h; vein R of male with small basal
patch of silver scales; vein Cu of male without silver scales. |
MALE GENITALIA. Sidepiece: Basal tergomesal area without dense patch of
long setae; median sternomesal area without strongly developed sclerite and with-
Out tuft; specialized subapical sternal seta absent. Prosophallus: Mesal lobe with
lateral portion usually inclined between 15° and 30° from horizontal; stem not
bowed; filament with hook not strongly angulate.
PUPA. Cephalothorax: Without pale inverted V-shaped marking; hair 5-C long,
more than twice length of 4-C; 9-C single. Abdomen: Hair 1-I with primary branches
predominantly at least double; 2-II laterad or mesad of 3-II, most often mesad for
less than 0.3 the distance from 1-II to 3-II, not more than 0.3; 3-III usually single.
50 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
Paddle: Pigmented, sometimes very weakly; apex not produced; hair 1-P long, sub-
equal in length to paddle.
LARVA. Head: Hairs 5,6-C multiple; 7-C more than half length of 6-C; 11-C
longer than mental plate; 14-C longer than mental plate, single or branched; bmh
branched. Antenna: Hair 1-A branched. Thorax: Hair 11-P less than half length of
14-P; 14-P usually branched; 3-M single; 4-M double to multiple; 8-T longer than
metathoracic pleural tubercle. Abdomen: Hair 5-I at least subequal in length to 4-I;
2-II well mesad of 4-II; 3-III double or triple; 10-III single; 3-VI branched; 4-VII
branched; 5-VII caudad of 4-VII. more than half length of 3-VII; 10,12-VII single.
Segment VIII: Midapical comb scale with free portion shorter than or subequal in
length to sessile portion; hair 2-VIII single or branched. Anal Segment: Saddle ex-
tending around segment for moderate distance, ventral margin without slit.
DISCUSSION. The monotypic Alboapicus Subgroup is characterized in the adults
by the silver scaled tarsi 5-II,I]I; in the pupa by the very long hairs 5-C and 1-P; and
in the larva by a combination of the multiple hair 5-C and the branched 1-A, 3-VI
and 4-VII. The male genitalia do not show distinctive subgroup characters.
This subgroup possibly represents an offshoot of the Terrens Subgroup (see Sys-
tematics and Evolution). It differs from the latter subgroup in the characters given
above and also in the tendency of hair 2-II of the pupa to be situated farther laterad.
The Alboapicus Subgroup occurs in the Canal Zone and in the lowlands of north-
eastern Panama.
9. Aedes (Finlaya) alboapicus Schick, n.sp.
Figs. 3,23,24
TYPE: Holotype ¢ (206A-15) with associated pupal and larval skins (5-114) and genitalia
slide (670912-1), Barro Colorado Island [elev. 100-500 ft], Canal Zone, Panama, 1 May 1945,
W.H.W. Komp [USNM]. Allotype @ (207B-28) with associated pupal and larval skins (43-94),
same data as holotype except collected 7 May 1943 [USNM]. Paratypes: 1 |p? (207B-14, 43-
130), 2 19 (207A-11, 43-90; 207A-31, 43-91), 2 2 (207B-44), same data as allotype [BM; UCLA].
Aedes alboapicus was a manuscript name of W.H.W. Komp for this species and is used here in
honor of Mr. Komp’s discovery. The name obviously refers to the silvered fifth tarsal segments.
FEMALE. Head: Vertex with decumbent scales laterad of median longitudinal
line dark, broad. Thorax (fig. 23): Prescutellar space and midlobe of scutellum
without silver scales. Legs: Femur I and usually femur II without posterior patch of
silver scales, femur II sometimes with sparse patch; tarsus 5-I sometimes sparsely sil-
ver scaled; tarsus 1-II with median dark band about 0.5-0.75; tarsus 5-II usually with
at least some silver scales, sometimes entirely silvered; femur III with basal dark band
0.09-0.14 (0.08-0.15), subapical dark band 0.27-0.30 (0.20-0.33); tarsus 1-III with
basal silver band 0.14-0.18 (0.13-0.20), apical silver band 0.15-0.19 (0.13-0.21).
Wing: Vein C with silver scales at base from small patch to line reaching crossvein h,
usually a short line; vein R without silver scales.
MALE. Head: Vertex with decumbent scales laterad of median longitudinal line
dark or silver, broad; palpus about 2 labellum lengths shorter than proboscis. Thorax
(fig. 23): Mesonotal disc with anterior silver band about 0.25-0.75 length of fossa
and with strip of silver scales bordering fossal macula; pra hairs pale or dark. Legs:
Tarsus 1-I] with median dark band about 0.5.
Schick: Terrens Group of Aedes (Finlaya) 51
MALE GENITALIA (fig. 23). Sidepiece: Length 0.32-0.33 mm (0.30-0.36 mm).
Prosophallus: Length 0.09-0.10 mm (0.09-0.11 mm); width 0.11-0.12 mm; mesal
lobe with lateral portion usually inclined between 15° and 30° from horizontal,
sometimes less than 15° or not appreciably inclined; stems usually divergent but
sometimes convergent; filament ratio 0.70-0.75. Aedeagus: Length 0.11-0.12 mm.
PUPA (fig. 23). Abdomen: Hair 9-VII long, subequal in length to paddle. Paddle:
Ventral midrib weakly pigmented throughout most length, not more strongly at
apex.
LARVA (fig. 24). Head: Hair 5-C usually 5-8 branched (4-8); 6-C triple to 5-
branched; 7-C usually 6,7-branched (5-8). Thorax: Hair 1-P usually triple to 5-
branched (3-7); 4-P usually 4-branched (2-7); 5-P usually double (2-4); 7-P usually
triple (2-4); 14-P usually double (1-3); 4-M usually double or triple (2-4). Abdomen:
Hair 3-VI usually double (2-3). Segment VIII: Comb scales 28 to about 60, scales
stout in 2 specimens with lowest number; length of free portion of midapical scale
0.024-0.028 mm; hair 2-VIII single or double. Siphon: Length 0.84-0.87 mm; L/S
2.48-2.86; P/L 0.49-0.53; H/L 0.58-0.59. Anal Segment: Ventral brush with 12
hairs (11-13); 4a-X usually 6,7-branched (5-7).
DISTRIBUTION (fig. 3). Canal Zone and Pacific lowlands of eastern Panama.
Material examined: 72 specimens; 8 6, 16 9, 17 pupae, 31 larvae; 24 individual rear-
ings (14 larval, 4 pupal, 1 incomplete). For the Komp collections see the explana-
tory chapter.
PANAMA. Canal Zone: Barro Colorado Island [elev. 100-500 ft], 7 May 1943, treehole, W.
Komp, type series in part, 2 Ip? (207B-14,18), 2 12 (207A-11,31), 2 2 (207B-44) [USNM; BM;
UCLA] — treehole, 1 Ipd (207B-27), 1 2 (207A-29), 2 1 (43-115,117; adult numbers unknown),
2 p? (207A-16), 1 1(43-121, adult number unknown) — treehole, 1 p? (207A-20) — treehole, 1 6,
1 2 (207B-29) — treehole, 1 1 (43-81, adult number unknown); 21 May 1943, very small treehole
in thin tree, 1 Ip? (207B-1), 1 p? (207B-23) — thin tree, 2 16 (207B-40,41), 1 19 (207A-9), 1°, 1
p, 2 1 (207A-34), 1 1 (207D-2) — treehole, 1 1 (207A-4), 2 1 (43-221, adult number unknown) —
small treehole, 1 Ipd (207A-7) — treehole, 1 9 (207A-10), 1 1 (43-160, adult number unknown) —
1 Ip (207A-24); 21 May 1943, 1 2;31 May 1943, 1 1d (207B-33); May 1943, treehole, W. Komp, 1
@ (207A-17) [UCLA; USNM]; 1 May 1945, W. Komp, 1 Ip¢ (206A-15), type series in part
[USNM]; 15 May 1945, small deep treehole, 1 p? (207C-13), 1 6 (207C-2), 1 2 (207C-7) — tree-
hole, 2 19 (207C-1,25) — 1 lp? (207C-6) — small treehole in thin stump, 1 lp? (207C-35) [UCLA;
USNM]. Darien: Morti, elev. 60 ft, 7 Dec 1966, treehole, height 1 ft, O. Berlin, R. Hinds, 1 lp?
(PA 989-10) [UCLA].
Buenaventura Subgroup
ADULTS. Head: Vertex of female with decumbent scales along longitudinal mid-
line apparently moderately broad, of male narrow or moderately broad; occiput of
both sexes usually with median erect scales pale and lateral erect scales dark; probos-
cis of female longer than femur I; palpal segment 3 of male with 2 to several short
hairs forming poorly developed tuft; segment 4 with hairs of ventrolateral row mod-
erately long, sparse. Thorax: Mesonotal disc of female not transversely silvered, of
male with anterior silver band; complete acrostichal or posterior dorsocentral lines
absent in female; acrostichal setae present; fossa macula of female not coextensive
with fossa, reduced mesally, of male coextensive with fossa; supraalar macula not
truncate posteriorly, in female broadly joined to fossal macula; ppn silver scaled; ssp
scales absent; pra hairs of female dark; upper stp and mep scale patches contiguous.
Legs: Mid- and hindlegs not shaggy; mid- and hindtarsi almost entirely dark scaled
52 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
(tarsus 1-II with inconspicuous narrow basal silver band); tarsus 1-I without apical
silver band; femur II without knee spot; femur III with basal dark band complete,
usually broad (incomplete in some males). Wing: Vein C of female with small basal
patch of silver scales, of male a line reaching crossvein h; vein R of male without sil-
ver scales or with small basal patch; vein Cu of male without silver scales.
MALE GENITALIA. Sidepiece: Basal tergomesal area with small patch of long
setae: median sternomesal area with strongly developed sclerite and with tuft; long
subapical sternal seta with apical twist present. Prosophallus: Mesal lobe with lateral
portion inclined at about 15° from horizontal; stem not bowed; filament with hook
not strongly angulate.
PUPA. Cephalothorax: Without pale inverted V-shaped marking; hair 5-C less than
twice length of 4-C; 9-C single. Abdomen: Hair 1-I with primary branches predom-
inantly double-triple to multiple; 2-II usually laterad of 3-II, when mesad, less than
0.3 the distance between 1-II and 3-II; 3-III single. Paddle: Clear or very weakly pig-
mented; apex not produced; hair 1-P shorter than paddle.
LARVA. Head: Hairs 5,6-C multiple; 7-C more than half length of 6-C; 11-C
longer than mental plate; 14-C longer than mental plate, branched; bmh branched.
Antenna: Hair 1-A single or branched. Thorax: Hair 11-P less than half length of 14-
P; 14-P branched; 3-M single; 4-M double; 8-T more than half length of but shorter
than metathoracic pleural tubercle. Abdomen: Hair 5-I at least subequal in length
to 4-I; 2-II well mesad of 4-II; 3-III double; 10-III single; 3-VI single; 4-VII branched;
5-VII cephalad of 4-VII, more than half length of 3-VII; 10-VII usually single; 12-
VII single. Segment VIIT: Midapical comb scale with free portion longer than sessile
portion; hair 2-VIII single. Anal Segment: Saddle extending around segment for
moderate distance, ventral margin without deep slit.
DISCUSSION. The monotypic Buenaventura Subgroup shows several unique char-
acters. In the adults tarsi 1-II,]II lack an apical silver band, tarsi 2-II,III lack a basal
silver band and the scale patches of the upper stp and mep are contiguous; in the
male genitalia a specialized subapical seta is present on the sidepiece and the aedea-
gus is more elongate than in the other species; and in the larva hair 5-VII is situated
cephalad of 4-VII. The pupa does not show distinctive subgroup characters.
Although possibly the most highly derived of the subgroups, Buenaventura shows
definite affinities to the subgroups that form the ‘“‘core’’ of the Terrens phyletic
line (see Systematics and Evolution). As in the Alboapicus Subgroup of this phyletic
line, the pupa differs from the Terrens Subgroup in the more lateral placement of
hair 2-IJ, but this condition is more extreme in the Buenaventura Subgroup. Whether
or not this is indicative of a common descent of the Alboapicus and Buenaventura
Subgroups is not known but the more lateral position of the hair possibly represents
the primitive condition.
The Buenaventura Subgroup occurs in the Pacific lowlands of Colombia.
10. Aedes (Finlaya) buenaventura Schick, n.sp.
Pigs. 3.25.26
TYPE: Holotype 3 (COL 56-102) with associated pupal skin and genitalia slide, Rio Raposo
Virus Field Station, ca 4 km W, nearest town Buenaventura, elev. 100 ft, Valle, Colombia, 23 Feb
1965, decaying trunk of Palma naidai, V.H. Lee [USNM]. Allotype ? with associated larval and
ee oe (56-11), same data as holotype [USNM]. Paratype: 1 L (56-1), same data as holotype
UCLA]. |
Schick: Terrens Group of Aedes (Finlaya) 53
FEMALE. Head: Vertex with decumbent scales laterad of median longitudinal
line dark, broad. Thorax (fig. 25): Most posterior acrostichal seta at about 0.33
from anterior end; prescutellar space and midlobe of scutellum without silver scales.
Legs: Femora I and II with posterior patch of silver scales in basal half well devel-
oped, about equally large; femur III with basal dark band 0.09-0.14, subapical dark
band 0.28-0.29, latter band extending to apex of segment along dorsal surface.
Wing: Vein R without silver scales.
MALE. Head: Vertex with decumbent scales laterad of median longitudinal line
silver, broad; palpus about 1 labellum length shorter than proboscis. Thorax (fig.
25): Mesonotal disc with anterior silver band about 0.25-0.5 length of fossa; acro-
stichal line usually absent; fossal macula coextensive with fossa. Legs: Similar to
those of female except that femur III with basal dark band sometimes incomplete
and scales of median silver band sometimes completely transecting subapical dark
band.
MALE GENITALIA (fig. 25). Sidepiece: Length 0.70 mm. Prosophallus: Length
0.11 mm; width 0.13 mm; stems convergent; filament ratio 0.70-0.75. Aedeagus:
Length 0.12 mm.
PUPA (fig. 25). Paddle: Ventral midrib clear or weakly pigmented for most
length.
LARVA (fig. 26). Head: Hair 5-C 5-7 branched; 6-C 6-9 branched; 7-C 7-9 branch-
ed. Thorax: Hair 1-P usually triple (2-4); 4-P usually triple (2-3); 5-P usually triple
(2-4); 7-P double. Abdomen: Hair 10-VII usually single (1-2). Segment VIII: Comb
scales more than 100; length of free portion of midapical scale 0.36-0.37 mm. Si-
phon: Length 0.53-0.56 mm; L/S 2.00-2.08; P/L 0.52-0.55: H/L 0.59-0.64. Anal
Segment: Ventral brush with 12 hairs; 4a-X 11-branched.
DISTRIBUTION (fig. 3). Pacific lowlands of department of Valle, Colombia. Ma-
terial examined: 27 specimens; 8 6, 3 , 11 pupae, 5 larvae; 11 individual rearings (4
larval, 7 pupal).
COLOMBIA. Valle: Rio Raposo Virus Field Station, ca 4 km W, nearest town Buenaventura,
elev. 100 ft, 23 Feb 1965, decaying trunk of Palma naidai, V. Lee (COL 56, type series), 1 lp? (56-
11), 2 pd (S6-100,102), 1 p? (56-101), 1 L (56-1) [USNM; UCLA]. Rio Raposo Virus Field Sta-
tion, ca 4 km N, elev. 70 ft, 5 May 1965, Palma naidai stump, P. Barreto, V. Rivas (COL 110), 2
Ipd (110-11,16), 1 lp? (110-17), 3 pd (100-12,100,108) [UCLA]. Rio Raposo Virus Field camp,
W of, in Rio Raposo brackish water zone, nearest town Buenaventura, elev. near sea level, 17 Dec
1965, large treehole, height ca 35 ft, V. Lee, 1 pd (COL 144-100) [UCLA].
Metoecopus Subgroup
ADULTS. Head: Vertex of both sexes with decumbent scales along longitudinal
midline narrow; occiput of both sexes with all erect scales dark; proboscis of female
longer than femur I; palpal segment 3 of male with long apical ventrolateral hairs
forming a tuft as long as segments 4 and 5 combined; segment 4 with hairs of ventro-
lateral row long, closely spaced. Thorax: Mesonotal disc of both sexes not trans-
versely silvered; complete acrostichal or posterior dorsocentral lines absent in fe-
male; acrostichal setae absent; fossal macula of both sexes coextensive with fossa, re-
duced mesally; supraalar macula not truncate posteriorly, in female broadly joined
to fossal macula; ppn silver scaled; ssp scales absent; pra hairs of female dark; upper
stp and mep scale patches not contiguous. Legs: Mid- and hindlegs not shaggy; mid-
and hindtarsi with conspicuous silver markings; tarsus 1-I with narrow apical silver
band; femur II with knee spot present, narrow to moderately broad, the scales not,
54 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
or at most, just reaching anterior subapical setae; tarsus 1-II with median dark band
complete, narrow to moderately broad; tarsus 2-II with complete apical dark band;
tarsus 5-II without silver scales; femur III with basal dark band complete, broad; tar-
sus 5-III without silver scales. Wing: Vein C of both sexes usually with small basal
patch of silver scales; vein R of males usually without silver scales; vein Cu of males
without silver scales.
MALE GENITALIA. Sidepiece: Basal tergomesal area without dense patch of
long setae; median sternomesal area without strongly developed sclerite and with-
out tuft; specialized subapical sternal seta absent. Prosophallus: Mesal lobe with
lateral portion usually inclined between 15° and 30° from horizontal; stem not
bowed; filament with hook not strongly angulate.
PUPA. Cephalothorax: Without pale inverted V-shaped marking; hair 5-C less than
twice length of 4-C; 9-C single. Abdomen: Hair 1-I with primary branches usually
predominantly single; 2-II mesad of 3-II for more than or less than 0.3 distance be-
tween 1-II and 3-II; 3-III single. Paddle: Pigmented; apex usually produced, often
strongly; hair 1-P shorter than paddle.
LARVA. Head: Hair 5-C usually single or double; 6-C usually single; 7-C more
than half length of 6-C; 11-C longer than mental plate; 14-C longer than mental
plate, usually single. Antenna: Hair 1-A single. Thorax: Hair 11-P less than half
length of 14-P; 14-P single; 3-M single; 4-M double; 8-T about subequal in length to
or longer than metathoracic pleural tubercle. Abdomen: Hair 5-I at least subequal in
length to 4-I; 2-II well mesad of 4-II; 3-III double; 10-III single; 3-VI single; 4-VII
single; 5-VII caudad of 4-VII, more than or less than half length of 3-VII; 10,12-VII
single. Segment VIII: Midapical comb scale with free portion longer than sessile por-
tion; hair 2-VIII single. Anal Segment: Saddle extending around segment for moder-
ate distance, ventral margin without deep slit.
DISCUSSION. The monotypic Metoecopus Subgroup is characterized in the fe-
male by the combination of the narrow lateral scales of the vertex, the broadly
joined fossal and supraalar maculae, the absence of ssp scales and the broad basal
band of femur III; and in the male by the dark erect scales of the vertex. The larva
can be distinguished from the previous subgroups by the single hair 4-VII but appar-
ently cannot be separated from those of species of the Homoeopus, Heteropus and
Podographicus Subgroups in which hairs 5,6-C are not multiple. The male genitalia
and pupa do not show distinctive subgroup characters.
As indicated in the chapter on Systematics and Evolution, the Metoecopus Sub-
group shows a tenuous relationship to the Terrens phyletic line, only 3 characters
indicating an affinity. The subgroup has little similarity with the only other sub-
group that occurs in the Pacific lowlands of Colombia, Buenaventura, aside from
the few Terrens phyletic line characters.
The subgroup occurs in the coastal lowlands of Ecuador.
11. Aedes (Finlaya) metoecopus Dyar
Figs. 3,27 ,28
1925. Aedes metoecopus Dyar, 1925a:30. TYPE: Lectotype 6 (2107/86) with genitalia slide
(2107), Ecuador, F. Campos Ribadeneira [USNM; designation of Stone and Knight 1956:
222}.
Schick: Terrens Group of Aedes (Finlaya) 55
Aedes (Finlaya) terrens metoecopus of Horsfall (1955:699).
Aedes (Finlaya) terrens var. metoecopus of Edwards (1932:150); Knight and Marks (1952:549).
Aedes (Finlaya) terrens in part of Lane (1939:105; 1953:686-687); Stone, Knight and Starcke
(1959:171); Belkin, Schick and Heinemann (1965:20); Forattini (1965:395).
Aedes (Gualteria) terrens in part of Levi-Castillo (1952a:555; 1952b:77).
Aedes (Finlaya) podographicus in part of Dyar (1928:223).
FEMALE. Head: Vertex with decumbent scales generally dark, narrow curved.
Thorax (fig. 27): Prescutellar space with silver scales; midlobe of scutellum with or
without silver scales. Legs: Femora I and II usually without posterior patch of silver
scales or, with small patch in basal half, infrequently with large but diffusely scaled
patch in basal half; tarsus 1-II with median dark band about 0.25-0.5; femur III with
basal dark band 0.11-0.17 (0.04 in 1 specimen), subapical dark band 0.36-0.41; tar-
sus 1-III with basal silver band 0.04-0.12, apical silver band 0.24-0.31 (0.21 in 1
specimen). Wing: Vein C without silver scales or with basal silver scales forming small
patch or line reaching about 0.5 to crossvein h; vein R without silver scales.
MALE. Head: Vertex with decumbent scales laterad of median longitudinal line
dark, broad; palpus about 3 labellum lengths shorter than proboscis. Thorax (fig.
27): Essentially as in female but fossal macula broader. Legs: Tarsus 1-II with med-
ian dark band about 0.25-0.5. Wing: Vein C usually with small basal patch of silver
scales and vein R without silver scales (in ECU 114-99 vein C with line of silver scales
reaching crossvein 4 and R with small basal patch of silver scales).
MALE GENITALIA (fig. 27). Sidepiece: Length 0.34-0.40 mm. Prosophallus:
Length usually 0.10-0.12 mm (0.13 mm unilaterally in 1 specimen); width 0.13-0.15
mm; mesal lobe with lateral portion inclined usually between 15° and 30° from hor-
izontal, sometimes at 15° or less; stems divergent; filament ratio 0.64-0.80. Aedea-
gus: Length 0.13-0.15 mm.
PUPA (fig. 27). Abdomen: Hair 1-I with primary branches usually predominantly
single, sometimes single-double or double; 2-II] mesad of 3-II for 0.1-0.5 the distance
from. 1-II to 3-II. Paddle: Ventral midrib strongly pigmented for entire length.
LARVA (fig. 28). Head: Hair 5-C usually single or double (1-3); 6-C usually single
(1-2); 7-C usually triple or 4-branched (2-5); 14-C usually single, rarely double. Thor-
ax: Hair 1-P usually triple (2-6); 4-P usually double to 4-branched (2-5); 5-P usually
double (1-3); 7-P usually double or triple, rarely single. Segment VIII: Comb scales
82-93; length of free portion of midapical scale 0.033-0.037 mm. Siphon: Length
0.80-1.02 mm; L/S 2.12-2.33 (2.00-2.38); P/L 0.57-0.60 (0.54-0.64); H/L 0.62-0.68.
Anal Segment: Ventral brush usually with 14 hairs (12-15); 4a-X usually 8-branched
6,8-10).
DISTRIBUTION (fig. 3). Lowlands of Guayas and Los Rios, Ecuador. Material
examined: 114 specimens; 17 6, 34 9, 25 pupae, 38 larvae; 25 individual rearings
(10 larval, 14 pupal, 1 incomplete).
ECUADOR. Guayas: Chongon (nearest town), Route 3, km 18-29, elev. 0-300 ft, 9 Feb 1966,
small treehole, J. Belkin, E. Gerberg (ECU 134), 1 lpd (134-10), 2 p? (134-100,101), 2 L (134-1)
[UCLA]. Empalme, 3 km S (ca 25 air miles NE Balzar), elev. 300-700 ft, 6 Feb 1966, biting-land-
ing, J. Belkin, et al, 1 9 (ECU 120-1) [UCLA]. Guayaquil [elev. near sea level] , Mar 1940, J. Mur-
dock (H-9-34), 1 6, 4 9; Apr 1940, J. Murdock (207B-24), 1 6, 1 2; Aug 1940, W. Komp, 1 2;
1943, 3 2(207B-19) [USNM] . Guayaquil, Country Club, J. Murdock (207A-45), 1 6, 1 @ [USNM].
Guayaquil (nearest town), Rt. 3, km 10, elev. 0-300 ft, 12 Feb 1966, cut or broken bam-
boo, J. Belkin, E. Gerberg, 1 L (ECU 161-3) [UCLA]. Pascuales, vicintiy of, Rt. 2,km 9.5, elev.
0-300 ft, 5 Feb 1966, small treeholes, height 6 ft, J. Belkin, W. Hjort, et al (ECU 105-107), 1 lp?
(107-10), 1 pd (105-100), 1 p? (107-100), 4 L (105-1), 2 L (106-2), 5 L (107-2) [UCLA] . Los Ri-
56 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
os: Hacienda Mora (near Montalvo), elev. 0-300 ft, 8 Feb 1966, cut bamboo, J. Belkin, E. Ger-
berg (ECU 125), 1 Ipé (125-11), 2 Ip? (125-12,14), 1 po (125-111), 4 L (125-1) [UCLA]. Valen-
cia, 4 km W, elev. 300-700 ft, 6 Feb 1966, fallen cacao pod, J. Belkin et al (ECU 114), 1 Ipd
(114-10), 4 Ip? (114-11-14), 4 pd (114-72,93,99), 5 p? (114-73,85 102,110,113), 1 IP (114-5), 9
L (114-4) [UCLA]. Department not specified: F. Campos R., 1 6 (139); 4 6; 1 2 (128); 1 2 (138);
1 2 (type series in part); 5 9 [USNM].
Insolitus Subgroup
ADULTS. Head: Vertex of both sexes with decumbent scales along longitudinal
midline narrow; occiput of both sexes with all erect scales pale; proboscis of females
longer than femur I; palpal segment 3 of males with several long apical ventrolateral
hairs forming a tuft as long as segments 4 and 5 combined; segment 4 with hairs of
ventrolateral row long, closely spaced. Thorax: Mesonotal disc of females not trans-
versely silvered, of males usually silvered for at least half length, sometimes not
transversely silvered; complete acrostichal or posterior dorsocentral lines absent in
females; acrostichal setae present or absent; fossal macula of females variable, coex-
tensive with fossa, variously reduced or absent, of males coextensive with fossa; su-
praalar macula not truncate posteriorly, in females broadly joined to fossal macula
(except when latter greatly reduced); ppn silver scaled; ssp scales present or absent;
pra hairs of females pale or dark; upper stp and mep scale patches not contiguous.
Legs: Mid- and hindlegs not shaggy; mid- and hindtarsi with conspicuous silver mark-
ings; tarsus 1-I with or without narrow apical silver band; femur II with knee spot us-
ually present, moderately broad to broad, the scales extending to level of or basad of
anterior subapical setae; tarsus 1-II with median dark band incomplete or complete,
narrow or moderately broad; tarsus 2-IJ with incomplete or complete dark apical
band or entirely silvered; tarsus 5-II usually without silver scales: femur III with bas-
al dark band complete, usually broad; tarsus 5-III without silver scales. Wing: Vein C
of females with basal line of silver scales usually reaching crossvein h, of males al-
ways reaching h; vein R of males with or without silver scales, when present, a
small basal patch or a line reaching level of crossvein h; vein Cu of males without
silver scales.
MALE GENITALIA. Sidepiece: Basal tergomesal area without dense patch of long
setae; median sternomesal area without strongly developed sclerite and without tuft:
specialized subapical sternal seta absent. Prosophallus: Mesal lobe with lateral por-
tion usually not appreciably inclined; stem not bowed; filament with hook not
strongly angulate.
PUPAE. Cephalothorax: With or without pale inverted V-shaped marking; hair 5-
C less than twice length of 4-C; 9-C single. Abdomen: Hair 1-I with primary branches
predominantly single to multiple; 2-II usually laterad of 3-II or mesad for less
than 0.3 the distance from 1-II to 3-II; 3-III usually single. Paddle: Usually clear or
very weakly pigmented; apex not produced; hair 1-P shorter than paddle.
LARVAE. Head: Hairs 5,6-C single to triple; 7-C more than half length of 6-C; 11-
C longer than mental plate; 14-C longer than mental plate, usually branched; bmh
usually branched. Antenna: Hair 1-A single. Thorax: 11-P greater or less than half
length of 14-P; 14-P single; 3-M single: 4-M double or triple; 8-T shorter, but more
than half length of, or longer than metathoracic pleural tubercle. Abdomen: Hair 5-I
at least subequal in length to 4-I; 2-II well mesad of 4-II; 3-III double or triple; 10-III
single or branched; 3-VI single or branched; 4-VII usually double; 5-VII caudad of 4-
Schick: Terrens Group of Aedes (Finlaya) 57
VII, more than half length of 3-VII; 10-VII single or branched; 12-VII single. Seg-
ment VIII: Midapical comb scale with free portion subequal in length to or longer
than sessile portion; hair 2-VIII single or branched. Anal Segment: Saddle extending
around segment for moderate distance, ventral margin without deep slit.
DISCUSSION. The Insolitus Subgroup is characterized primarily by the males.
The mesonotal disc is usually largely silvered, as in the Homoeopus Subgroup, but
unlike in the latter the basal dark band of femur III is complete and broad and vein
R is not as extensively silvered. The females show a fossal macula that is variously re-
duced and sometimes absent but otherwise cannot be separated from those of the
Terrens Subgroup. The larvae are characterized by a combination of the usually
branched but non-multiple hairs 5,6-C, the branched 4-VII and the free portion of
the midapical comb scale which is longer than or subequal in length to the sessile
portion. The male genitalia and pupa do not show distinctive subgroup characters.
The Insolitus Subgroup, as suggested in the chapter on Systematics and Evolution,
represents the most primitive of the subgroups of the Terrens Group and the stock
ancestral to the other subgroups. It almost certainly belongs to the Terrens phyletic
line but differs from the other subgroups of the line in the usually extensively sil-
vered mesonotum of the males and its distribution.
The subgroup occurs in the highlands of Central America (Guatemala, Panama),
on the Atlantic coast of Panama, in the highlands of Colombia and northern Vene-
zuela and in Trinidad.
Only | species, insolitus, is included in the subgroup but a second species is now
known from the Rancho Grande area of Venezuela (see insolitus). The subgroup
description is based upon both species.
12. Aedes (Finlaya) insolitus (Coquillett)
Figs. 4,29 30
1906. Verrallina insolita Coquillett, 1906a:62. TYPE: Holotype ?, Trinidad, F.W. Urich [USNM,
9142].
1906. ‘Verrallina laternaria Coquillett, 1906b:184. TYPE: Holotype 3, Montserrat, Caroni, Trin-
idad, June 1905, August Busck [USNM, 8290]. Type data from Belkin, Schick and Hei-
nemann (1965:69). NEW SYNONYMY.
Aedes insolitus of Dyar (1906:16-17).
Verrallina insolita of Coquillett (1906c:17); Theobald (1910:496); Surcouf and Gonzales-Rincon-
es (1911-226),
Aedes (Finlaya) terrens var. podographicus in part of Edwards (1932:150); Knight and Marks
(1952:549).
? Aedes podographicus of Boshell-Manrique and Osorno-Mesa (1944: 173).
Aedes (Finlaya) terrens in part of Dyar (1921:152); Bonne and Bonne-Wepster (1925 :424); Dyar
(1928:224); Lane (1939:105; 1953:686-687); Stone, Knight and Starcke (1959:171); Belkin,
Schick and Heinemann (1965:69); Forattini (1965 :395).
Aedes oswaldi in part of Urich (1913:528); Howard, Dyar and Knab (1917:815-819).
Aedes laternaria of Dyar (1906:17); Dyar and Knab (1906b:191,202).
Verrallina laternaria of Coquillett (1906c:17); Theobald (1910:496).
FEMALE. Head: Vertex with decumbent scales laterad of median longitudinal
line broad, usually all dark. Thorax (fig. 29): Mesonotal disc, in forms with rela-
58 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
tively well-developed fossal macula (see below) often with a short anterior silver
acrostichal line, and in forms with reduced macula without line; acrostichal setae
variously developed [always present in Trinidad, most posterior seta usually at
about 0.33 from anterior end, sometimes as far caudad as about 0.5; usually ab-
sent in other populations, when present in these, most posterior seta at about
0.25]; fossal macula extremely variable in development, infrequently coextensive
with fossa (fig. 29, TR 827-114), usually variously reduced mesally and central-
ly (e.g. fig. 29, TR 778-117) or rarely absent; prescutellar space usually without sil-
ver scales; midlobe of scutellum with silver scales; ssp scales present; pra hairs pale.
Legs: Femur I with well-developed posterior patch of silver scales in basal half; fe-
mur II in Central American highlands and Trinidad with posterior patch usually larg-
er than that of femur I but in coastal Central America patch smaller than that of fe-
mur I; femur II with knee spot usually present (absent in TR 680-105); tarsus 1-II
with median dark band complete in Central America, about 0.33-0.5, in other local-
ities usually incomplete and sometimes entirely obliterated by silver scales, when
complete 0.25-0.33; tarsus 2-II in most localities with complete dark apical band, in
Trinidad often entirely silver scaled or with at least some apical silver scales; tarsus
5-II usually dark scaled (silver scaled in 1 specimen of TR 620); femur III with
basal dark band 0.06-0.15 [in San Felipe, Guatemala, 0.06; in Cerro Mali, Panama,
0.08-0.11; in other localities 0.11-0.14 (0.10-0.15)], subapical dark band 0.21-0.33
[in most localities 0.21-0.27 (0.20-0.28); in Colombia 0.27-0.33]; tarsus 1-III with
basal silver band absent or 0.04-0.15 [in most localities 0.04-0.09 (absent-0.12); in
coastal Central America 0.08-0.15], apical silver band 0.12-0.24 (0.12-0.30; entire
range in Trinidad). Wing: Vein R usually without silver scales, sometimes with a few
at base.
MALE. Head: Vertex with decumbent scales laterad of median longitudinal line
silver, broad; palpus in most localities about 1-2 labellum lengths shorter than pro-
boscis, in Isabi, Colombia, subequal in length to proboscis. Thorax (fig. 29): Meso-
notal disc in most localities almost entirely silver scaled (fig. 29, TR 782-105), in
Trinidad dorsocentral area silver or dark scaled, in latter case acrostichal line per-
sisting as more or less prominent marking (fig. 29, TR 827-109); pleuron as in fe-
male. Legs: Essentially as in female. Wing: Vein R with basal silver marking variously
developed [in Volcanes, Colombia, absent; in Trinidad usually absent, infrequently
a small basal patch; in Capira, Panama, absent or a small basal patch (2 specimens);
in Yepocapa, Guatemala and Isabi, Colombia, a small basal patch and also some
silver scales disjunctly at level of crossvein h, in San Felipe, Guatemala, and in
Cuincha and Muzo, Colombia, a line reaching level of h].
MALE GENITALIA (fig. 29). Sidepiece: Length 0.54-0.70 mm. Prosophallus:
Length 0.08-0.09 mm (0.07-0.10 mm); width 0.11-0.12 (0.11-0.13 mm); mesal lobe
with lateral portion usually not appreciably inclined or inclined at less than 15° from
horizontal, rarely as great as between 15° and 30°; stems divergent in most localit-
ies (convergent in Cerro Mali and Cerro Campana, Panama); filament ratio 0.85-1.20
(0.75-1.35). Aedeagus: Length 0.11-0.13 mm.
PUPA (fig. 29). Cephalothorax: Pale inverted V-shaped marking not developed or
weakly differentiated. Abdomen: Hair 2-II laterad or mesad of 3-II [in Cerro Mali,
Panama, laterad of 3-II to mesad for less than 0.3 the distance from 1-II to 3-Il:
in Trinidad infrequently laterad, usually mesad from less than 0.3 to more than 0.3:
in other localities placement of 2-II intermediate between above. conditions] ; 3-III
usually single, rarely branched. Paddle: Showing geographic dimorphism as follows,
but intermediate types present in Trinidad. Central American type (fig. 29, PA
Schick: Terrens Group of Aedes (Finlaya) 59
497): Clear or very weakly pigmented; not tapered; apex not produced, often emar-
ginate; ventral midrib not or weakly pigmented. Trinidad type (fig. 29, TR 778):
Pigmented; tapered; apex also not produced but not emarginate; ventral midrib mod-
erately pigmented, not more strongly at apex.
LARVA (fig. 30). “‘Northern highlands” in this description refers to San Felipe,
Guatemala (GUA 130) and Cerro Campana, Panama (PA 497). Head: Hair 4-C single
to triple [in Northern highlands 1(1-2); in Trinidad 1,2(1-3); in Almirante, Panama,
2; in Cerro Mali, Panama, 2(1-3)] ; 6-C single to triple [in Northern highlands 1(1-2);
in Almirante 3; in Trinidad 3(1-3); in Cerro Mali 3(2-3, rarely 1)]; 7-C double to 7-
branched [in Northern highlands 2,3(2-4); in Cerro Mali 3-5(2-7); in Trinidad 4,5
(2-6); in Almirante 6,7]; 11-C in Cerro Campana about 1.5 length of mental plate,
in other localities at least twice length of plate; 14-C in Northern highlands finer and
tending to be shorter than in other localities, usually double (2-3, rarely 1); bmh us-
ually double (2; rarely 1,3). Thorax: Hair 1-P single to 5-branched [in Northern high-
lands 2(1-2); in other localities 3,4(2-5)]; 4-P single to 4-branched [in Cerro Cam-
pana 1; in San Felipe 1-3; in Almirante 2; in Trinidad 2(2-3, rarely 1); in Cerro Mali
3(2-3, rarely 4)]; 7-P single to 4-branched [in Northern highlands 1; in Almirante 2;
in Trinidad 2(1-3);in Cerro Mali 3(2-4)]; 11-P variable in length [less than half length
of 14-P in Almirante and Cerro Campana, Panama; at least half length of 14-P in
other localities]; 14-P single; 8-T shorter or longer than metathoracic pleural tubercle,
particularly short in Cerro Campana. Abdomen: Hair 5-I usually subequal in length
to or longer than 4-I, sometimes shorter than 4-I in Cerro Campana; 7-III sometimes
long, subequal in length to 7-II; 3-III usually double (2-3); 10-III single or double; 3-
VI single or double; 4-VII usually double (1-2); 10-VII usually single, sometimes
double in Trinidad. Segment VIIT: Comb scales 40 to more than 100; length of free
portion of apical scale 0.027-0.052 mm [in San Felipe 0.027-0.033 mm; in Cerro
Campana 0.033-0.035 mm; in Almirante 0.034-0.035 mm; in Trinidad 0.036-0.047
mm (0.035-0.048 mm, 0.027 mm in 1 specimen with atypical broad ligulate rather
than pandurate scale)]. Siphon: Length 0.61-0.92 mm [in Northern highlands 0.61-
O.72 mm; in Cero Mali 0.77-0.82 (0.72-0.87 mm); in Trinidad 0.67-0.87 mm (0.61-
0.92 mm); Almirante unknown]; L/S 2.12-2.45 [in most localities 2.22-2.38 (2.12-
2.42); in Northern highlands 2.27-2.46]; P/L 0.49-0.58 [in Cerro Mali 0.49-0.52
(0.47-0.55); in Northern highlands 0.49-0.54; in Trinidad 0.51-0.55 (0.48-0.58)];
H/L 0.52-0.65 [in Cerro Mali 0.56-0.59 (0.52-0.59); in Northern highlands 0.54-
0.61; in Trinidad 0.60-0.63 (0.57-0.65)]. Anal Segment: Ventral brush with 12 hairs,
(11-13); 4a-X usually 7-branched (5-8).
SYSTEMATICS. Aedes insolitus is widely distributed and shows some marked
geographical variation and perhaps actually represents a species complex. Some re-
gional populations are briefly discussed below.
In San Felipe, Guatemala, and Cerro Campana, Panama (‘‘Northern highlands’’),
certain of the larval hairs tend to have fewer branches than in other localities. Hair
6-C is single rather than usually triple, 1-P usually double rather than triple or mult-
iple and 5,7-P is single rather than usually double or triple. Also, 14-C is finer and
tends to be shorter than in other localities and the siphon is shorter than elsewhere,
0.61-0.72 mm vs. 0.77-0.92 mm.
The topotypic populations of Trinidad show variation in 2 characters that are
not variable in other populations, (1) the dorsocentral area of mesonotal disc of the
male is either largely silver scaled or dark scaled (fig. 29) and (2) the paddle of
the pupa is either clear and broadly rounded (and sometimes emarginate at apex)
or moderately well pigmented and somewhat tapered, with intermediate paddle
60 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
types often being developed. This first mentioned state of the above characters is
the condition shown by the other populations (the immature stages of insolitus
from Colombia are unknown) and the second mentioned alternative the condition
shown only by the Trinidad populations. There is a strong correlation of the non-
Trinidad vs. the Trinidad states and | or the other predominates or is exclusively
represented in the various lots. Two further differences between non-Trinidad and
Trinidad populations are (1) acrostichal setae are usually absent in the former and
present in the latter and (2) hair 2-II of the pupa tends to be situated farther mesad
in the latter.
The significance of the variation in Trinidad is unknown. Perhaps 2 species are
involved or perhaps hybridization has occurred between an ancestral population
showing the Trinidad character states and a species showing the non-Trinidad states.
The male genitalia and larvae-do not show corresponding variation.
Males from Isabi, Colombia, show an unusual character for the species, a very
long palpus in the male.
A second species of the Insolitus Subgroup came to my attention too late to be
formally included in the present revision. This species, from the Rancho Grande
area of Venezuela, can be distinguished from insolitus by the absence of ssp scales
and the dark pra hairs. The dorsocentral silvering of the males extends caudad into
the posterior half of the disc. |
DISTRIBUTION (fig. 4). Central America and Colombia at elevations of 2600-
5000 ft; Atlantic lowlands of Costa Rica and Panama; Trinidad at elevations of 200-
3000 ft. Material examined: 487 specimens; 58 6, 86 2, 106 pupae, 217 larvae; 89
individual rearings (58 larval, 23 pupal, 13 incomplete).
COLOMBIA. Boyaca: Cuincha (near Muzo), [elev. 1600-3300 ft], 17 Mar 1943, E. Osorno-
Mesa, 1 6 [UCLA]. Isabi (near Muzo), [elev. 3300-5000 ft], 18 Apr and 6 May 1942, E. Osorno-
Mesa, 3 6 [UCLA]. Muzo [elev. 1600-3300 ft] , 28 May 1963, E. Osorno-Mesa, 2 ¢ (161) [UCLA].
Cundinimarca: Volcanes, a patch of forest near Caparrapi, elev. 3300-5000 ft, 19 Sept 1943, H.
Kumm, | 6 (13), 1 2 (11); 1943, Kumm, 21 [UCLA].
COSTA RICA. Limon: Suerre, 20 July 1923, A. Alfaro, 3 6,5 9 [USNM].
GUATEMALA. Chimaltenango: Locality not specified, nearest town Yepocapa [elev. 4850
ft], 27 July 1950, treehole, P. Garcia, M. Xinic, 2 9 (GUA 140-1): 29 July 1950, treehole, H.
Dalmat, 2 d (GUA 141-1) [UCLA]. Suchitepequez: Cafetal Hamburgo (near San Felipe), elev.
ca 2600 ft, 8 Sept 1964, cut or broken bamboo, Almengar and P. Cowsill (GUA 130), 2 Ipd
(130-11,14), 2 Ip? (130-12,13), 3 pd (130-104-106), 1 p? (130-103) [UCLA]. Department not
specified: 1 6 (GUA 151-36) [UCLA].
PANAMA. Bocas del Toro: Almirante, elev. ca 100 ft, 30 Apr 1963, treehole (PA 290), 2 pd
(290-101,103), 1 p? (290-105), 1 p, 2 1(290-1) [UCLA] . Darien: Cerro Mali, western slope, elev.
4650 ft, 28 May 1963, treehole in small tree, height 7 ft (PA 368), 3 lpd (368-103,128,140), 15
Ip? (368-108 ,109,116-119,123,133-139,141), 6 pd (368-101,105,107,112,114,115), 5 p? (368-
102,104,106,113,136), 9 IP (368-110,111,120,122,124,126,127,131,132),4 P, 114 L, 1 1(368-1)
[UCLA]. Cerro Mali, auxiliary ridge W of, elev. 4900 ft, 22 May 1963, treehole, height 20 ft (PA
349), 1 Ip? (349-103), 1 L (349-1) [UCLA] . Panama: Cerro Campana, elev. 3500 ft, 12 Aug 1963,
treehole, height 3 ft (PA 497), 3 lpé (497-105-107), 1 lp? (497-101), 2 pd (497-102,103), 14 L
(497-1) [UCLA].
TRINIDAD. Mayaro: Bush Bush Forest, 16 Oct 1964, biting, 900-1300 hrs, height O-3 ft, Trul,
1 2? (TR 773-101) {UCLA]. St. Andrew: Cumaca, elev. 250-500 ft, 22 Oct 1964, large treehole,
height 10 tt A. Guerra CI K 776,779), 5 lod (7 78-1151 16,125; 799-122 130), 1 16 (779-110), 13
Ip? (778-117-121 123,124,127; 779-118-121 ,135), 2 pd (779-115,116), 1 IP (779-137), 6 L (778-
1), 2 6, 4 L (779-1) — large tub (TR 782), 1 Ipé (782-111), 2 Ip? (782-112,121), 1 pd (782-105),
1 Ip (782-110), 1 L (782-5); 14 Jan 1965, large treehole, height 15 ft, A. Guerra (TR 932), 1 lpd
(932-121), 1 Ip? (932-122) — large tub, 1 lp? (TR 940-127); 18 Feb 1965, large tub, A. Guerra
Schick: Terrens Group of Aedes (Finlaya) 61
(TR 1009), 2 Ip? (1009-40,42), 1 L (1009-4) [UCLA]. Mt. Tamana, elev. 1000 ft, 23 Aug 1964,
biting, 1100-1300 hrs, height 1-3 ft, R. Manuel and R. Martinez, 3 ? (TR 620) [UCLA]. St.
George: El Tucuche [elev. 3000 ft] , rain barrel, June 1942, 1 9; rain barrel, 2 d (29V142-2, 20VII-
42-1) [USNM]. Las Lapas Trace, elev. 2000 ft, 3 Apr 1964, biting, 1000-1500 hrs, height 4 ft, A.
Guerra (TR 281), 6 9 [UCLA]. Mt. Beck, elev. 800 ft, 29 Apr 1965, biting, 1400 hrs, height 3 ft,
A. Guerra, 1 ? (TR 1143-2) [UCLA]. Verdant Vale, 4 mi on Blanchisseuse road, nearest town
Arima, elev. 500 ft, 12 Nov 1964, large treehole, height 2 ft, A. Guerra (TR 827), 5 Ipd (827-
109-112,118), 3 Ip? (827-113,114,117), 1 p? (827-117), 2 2, 2 p (827-1) — cut bamboo (TR
829), 1 lp? (829-115), 1 IP (829-119) — 1 Ip? (TR 830-101) [UCLA]. Verdant Vale; elev. 1000
ft, large treehole, height 8 ft, A. Guerra (TR 680), 1 Ip? (680-105), 1 IP (680-104), 1 L (680-1)
[UCLA] . Locality not specified: F. Amandes, 14 Nov 1965, F. Urich, 1 2; 18 Nov 1905, F. Urich,
2d. 1 Si June, A. Busck; 1 6434.2 9d); 26,4 2) F. Urich’ 3 O(27-7 8.10). 6 927-911 ,12;, 812-7,
B-13-8,12) [USNM].
Aitkeni Subgroup
ADULT. (Female only.) Head: Vertex with decumbent scales along median lon-
gitudinal line narrow; occiput with median erect scales pale, lateral scales dark;
proboscis longer than femur I. Thorax: Mesonotal disc with anterior silver band;
complete acrostichal or posterior dorsocentral lines absent; acrostichal setae pres-
ent; fossal macula not coextensive with fossa, reduced mesally; supraalar macula not
truncate posteriorly, broadly joined to fossal macula; ppn dark scaled; ssp scales
present; pra hairs pale; upper stp and mep scale patches not contiguous. Legs: Mid-
and hindlegs not shaggy; mid- and hindtarsi with conspicuous silver markings; tarsus
1-I without apical silver band; femur II with knee spot broad, the scales extending
basad of anterior subapical setae; tarsus 1-I] with median dark band complete, broad;
tarsus 2-II with complete apical dark band; femur III with basal dark band incom-
plete; tarsus 5-III without silver scales. Wing: Vein C without silver scales.
PUPA. Cephalothorax: Without pale inverted V-shaped marking; hair 5-C less than
twice length of 4-C; 9-C single. Abdomen: Hair 1-I with primary branches predom-
inantly single-double; 2-II laterad of 3-II; 3-III single. Paddle: Pigmented; apex not
produced; hair 1-P shorter than paddle.
LARVA. Unknown.
DISCUSSION. The monotypic Aitkeni Subgroup is known only by the adult fe-
male and pupa. The female shows 2 unique characters for the Terrens Group, the
presence of pale gold scales in the fossal macula and the presence of dark rather
than silver scales on the ppn. The pupa does not show distinctive subgroup char-
acters.
The subgroup occurs at high elevations in southern Central America. Its phyletic
affinities are obscure but it possibly represents an early offshoot of the ancestral
stock of the Terrens Group.
13. Aedes (Finlaya) aitkeni Schick, n.sp.
Figs. 4,39
TYPE: Holotype ? (CR 50-101) with associated pupal skin, La Sierra, 10 km SE on hwy 2,
elev. 8000 ft, San Jose, Costa Rica, 24 Nov 1962, rotted depression in fallen tree, C.L. Hogue and
W.A. Powder [USNM]. Paratype: 1 p (50-2), same data as holotype [UCLA] . This species is dedi-
cated to T.H.G. Aitken.
62 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
FEMALE. Head: Vertex with broad median longitudinal line of narrow curved
scales; decumbent scales laterad of line dark, narrow curved. Thorax (fig. 39): Meso-
notal disc with narrow anterior band and a short silver acrostichal stripe extending
caudad from band; most posterior acrostichal seta at about 0.5 from anterior end of
acrostichal line; prescutellar space without silver scales; midlobe of scutellum with
silver scales. Legs: Femora I and II with well-developed posterior patch of silver
scales in basal half, that of femur II larger; femur III with basal dark band incom-
plete, subapical dark band 0.33; tarsus 1-IIJ with basal and apical dark bands each
0.08. Wing: Vein R without silver scales.
MALE. Unknown.
PUPA (fig. 39). Paddle: Weakly emarginate; ventral midrib moderately pigmented
for most length, weakly at apex.
LARVA. Unknown.
SYSTEMATICS. Pedro Galindo saw the holotype specimen of aitkeni during a
visit to the laboratory here at UCLA and believes it is the same form he has found
at high elevation in the province of Chiriqui, Panama, that fiercely bites man. The
Panamanian record cited for this species is based upon Galindo’s identification.
DISTRIBUTION (fig. 4). Province of San Jose, Costa Rica, elevation 8000 ft; pos-
sibly in Chirique highlands of Panama (Galindo, personal communication). Material
examined: 3 specimens; 1 9, 2 pupae; | individual rearing (pupal).
COSTA RICA. San Jose: La Sierra, 10 km SE on hwy 2, elev. 8000 ft, 24 Nov 1962, rotted
depression in fallen tree, C. Hogue, W. Powder (CR SO, type series), 1 p? (50-101), 1 p (50-2)
[UCLA].
Homoeopus Subgroup
ADULTS. (Female known for only 1 species.) Head: Vertex of both sexes with
decumbent scales along longitudinal midline narrow; occiput of female with erect
scales predominantly pale, of males all pale; proboscis of female longer than femur I;
palpal segment 3 of males with several long apical ventrolateral hairs forming a tuft
subequal in length to segments 4 and 5 combined; segment 4 with hairs of ventro-
lateral row long, somewhat sparse to closely spaced. Thorax: Mesonotal disc of
female not transversely silvered, of males usually transversely silvered for nearly en-
tire length; complete acrostichal or posterior dorsocentral stripes absent in female;
acrostichal setae present; fossal macula of female not coextensive with fossa, reduced
mesally, of males coextensive with fossa; supraalar macula not truncate posteriorly,
in females not joined to or narrowly or broadly joined to fossal macula; ppn silver
scaled; ssp scales present; pra hairs of female pale; upper stp and mep scale patches
not contiguous. Legs: Mid- and hindlegs not shaggy; mid- and hindtarsi with conspic-
uous silver markings; tarsus 1-I with or without narrow apical silver band; femur II
with knee spot present, broad, the scales extending basad of anterior subapical setae;
tarsus 1-I] with median dark band incomplete or complete and moderately broad;
tarsus 2-II with complete apical dark band; tarsus 5-II without silver scales; femur III
with basal dark band usually incomplete, narrow when complete; tarsus 5-III with-
out silver scales. Wing: Vein C of female with small patch of silver scales or line not
reaching 0.5 to crossvein h, of males reaching h; vein R of males with line of basal
silver scales extending well beyond level of crossvein h (about twice length of line
on C); vein Cu of males with basal silver scales.
MALE GENITALIA. Sidepiece: Basal tergomesal area without dense patch of long
Schick: Terrens Group of Aedes (Finlaya) 63
setae; median sternomesal area with or without strongly developed sclerite, with or
without tuft; specialized subapical sternal seta absent. Prosophallus: Mesal lobe with
lateral portion usually inclined between 15° and 30° from horizontal; stem usually
not bowed; filament with hook strongly or not strongly angulate.
PUPAE. Cephalothorax: Without pale inverted V-shaped marking; hair 5-C less
than twice length of 4-C; 9-C usually double. Abdomen: Hair 1-I with primary
branches usually predominantly single or single-double; 2-II usually laterad of 3-II
or mesad for 0.3 or less the distance from 1-II to 3-II; 3-III single. Paddle: Pig-
mented; apex not or weakly produced; hair 1-P shorter than paddle.
LARVAE. Head: Hair 5-C single to triple; 6-C single to double; 7-C more than
half length of 6-C; 11-C longer than mental plate; 14-C longer than mental plate,
single; bmh single. Antenna: Hair 1-A single. Thorax: Hair 11-P less than half length
of 14-P; 14-P single; 3-M single; 4-M usually double; 8-T shorter, but more than
half length of, or longer than metathoracic pleural tubercle. Abdomen: Hair 5-I at
least subequal in length to 4-I; 2-II well mesad of 4-II; 3-III] usually double; 10-
III single; 3-VI single; 4-VII usually single; 5-VII caudad of 4-VII, more than or less
than half length of 3-VII; 10,12-VII single. Segment VIIT: Midapical comb scale with
free portion longer or shorter than sessile portion; hair 2-VIII single. Anal Segment:
Saddle extending around segment for moderate distance; ventral margin without
deep slit.
DISCUSSION. The Homoeopus Subgroup is characterized primarily in the males;
the mesonotal disc is usually largely silvered, vein R is very extensively silvered, the
scales forming a basal line extending well beyond the level of crossvein h, and vein
Cu is basally silvered. The female of only homoeopus is known and it does not dif-
fer from the females of the Heteropus Subgroup in any fundamental respects. In
the larvae hairs 5,6-C are not multiple and 4-VII is single. I have been unable to
separate these from the larvae of the species in the Metoecopus, Heteropus and
Podographicus Subgroups that show non-multiple head hairs. The male genitalia
and pupae do not show distinctive subgroup characters.
Two species, homoeopus and impostor, comprise the Homoeopus Subgroup. The
former, although widely distributed, appears to be a relatively old form since it is
represented in the state of Veracruz, Mexico, by a relict population in which the
females show the primitive character of broadly joined fossal and supraalar macu-
lae. Aedes impostor appears to be an offshoot of homoeopus, considering the ap-
parent antiquity of homoeopus and that the genitalia of impostor possibly represent
a simplification of the homoeopus type (the male genitalia of homoeopus shows
variation in which the impostor condition is approached).
The subgroup occurs at moderately high elevations in the state of Veracruz, Mex-
ico, and in Central America from Guatemala southward into Costa Rica.
14. Aedes (Finlaya) homoeopus Dyar
Figs. 4,31,32
1922. Aedes terrens homoeopus, Dyar 1922:92-93. TYPE: Lectotype 6, Alajuela, Costa Rica,
Oct 1921, A. Alfaro [USNM; designation of Stone and Knight 1956:219].
Aedes (Finlaya) homoeopus of Bonne and Bonne-Wepster (1925:424-428).
Aedes (Finlaya) argyrothorax in part of Lane (1951:335; 1953:688); Stone, Knight and Starcke
(1959:159); Belkin, Schick and Heinemann (1965:12); Forattini (1965 :389).
64 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
Aedes (Finlaya) terrens var. podographicus of Edwards (1932:150); Knight and Marks (1952:549).
Aedes (Finlaya) terrens in part of Dyar (1928:224); Lane (1939:105).
Aedes terrens in part of Kumm, Komp and Ruiz (1940:400,417); Kumm and Zuniga (1942:404);
Vargas (1949a:221).
Aedes oswaldi in part of Howard, Dyar and Knab (1918:819).
FEMALE. Head: Vertex with decumbent scales laterad of median longitudinal
line dark, narrow curved. Thorax (fig. 31): Acrostichal and posterior dorsocentral
lines usually absent, sometimes weakly developed; fossal macula usually moderately
well developed, rarely reduced forming narrow diffusely scaled stripe; supraalar mac-
ula in Central America usually narrowly joined to fossal macula but sometimes not
joining macula, in Mexico broadly joined to macula; prescutellar space with sil-
ver scales; midlobe of scutellum usually with silver scales. Legs: Femora I and II
with well-developed posterior patch of silver scales in basal half, that of femur III lar-
ger; tarsus 1-II with median dark band in most localities about 0.4-0.5, in Costa Rica
about 0.33-0.5; femur III with basal dark band usually incomplete, when complete
at most 0.07, subapical dark band 0.32-0.37 (0.29-0.39); tarsus 1-III with basal silver
band 0.12-0.16 (0.11-0.18), apical silver band 0.17-0.22 (0.15-0.26). Wing: Vein R
without silver scales or with small basal patch, former condition common in Guate-
mala but infrequent elsewhere. :
MALE. Head: Vertex with all decumbent scales silver, narrow curved; palpus
about 0.5-2 labellum lengths shorter than proboscis, segment 4 with hairs of ven-
trolateral row somewhat sparse to closely spaced, latter condition most frequent
in Mexico and Guatemala. Thorax (fig. 31): Mesonotal disc usually transversely sil-
vered for at least half length, infrequently more narrowly, rarely with no transverse
silvering (GUA 48-11; this specimen with poorly defined strip of silver scales bord-
ering fossal macula and with complete acrostichal line). Legs: Tarsus 1-II with
median dark band variously developed [in El Salvador incomplete; in Mexico and
Costa Rica complete, about 0.33-0.5; in Guatemala complete, about 0.5 orgreater|.
Wing: Vein Cu usually with at least some silver scales on stem, often forming long
line:
MALE GENITALIA (fig. 31). Sidepiece: Length 0.34-0.41 mm [in Costa Rica
0.34-0.37 mm; in El Salvador 0.35-0.38 mm; in Guatemala 0.36-0.40 mm; in Mexico
0.40-0.41 mm] ; median sternomesal area with convexity usually strongly developed,
sometimes weakly or moderately so, sclerite typically well developed, tuft present.
Prosophallus: Length 0.11-0.14 mm [in El Salvador 0.11 mm; in Costa Rica 0.1 1-
0.13 mm; in Guatemala 0.11-0.14 mm; in Mexico 0.13 mm]; mesal lobe with lateral
portion usually inclined between 15° and 30° from horizontal (at about 45° in 1
specimen); filament ratio 0.55-0.75 (0.40-0.85), hook usually strongly angulate (not
strongly angulate in 2 specimens from Guatemala). Aedeagus: Length 0.13-0.14
mm (0.12-0.14 mm).
PUPA (fig. 31). Cephalothorax: Hair 9-C usually double, sometimes single. A bdo-
men: Hair 1-I with primary branches usually predominantly single or single-double,
sometimes predominantly double or double-triple; 2-I] laterad of 3-II to mesad
for more than 0.3 the distance from 1-II to 3-II, usually in line with 3-II or mesad
for 0.3 or less, rarely mesad for more than 0.3. Paddle: Ventral midrib with pigmen-
tation moderate to strong and usually extending to apex. |
LARVA (fig. 32). Head: Hair 5-C single to triple [in Amatitlan, Guatemala, single;
in Costa Rica usually single (1-2, rarely 3); in Mexico triple]; 6-C single or double
[in Amatitlan single; in Costa Rica usually single (1-2); in Mexico single or double];
Schick: Terrens Group of Aedes (Finlaya) 65
7-C double to 4-branched; 14-C usually single, rarely double; bmh usually single,
rarely double. Thorax: Hair 1-P double to 4-branched [in Amatitlan triple; in Costa
Rica usually triple (2-4); in Mexico 4-branched (1 specimen)]; 4-P double to 6-
branched [in Costa Rica usually triple (2-4); in Amatitlan 4-branched; in Mexico 3,4,
6-branched]; 5-P single to triple [in Amatitlan usually double (1-2); in Costa Rica
usually double (2-3, rarely 1); in Mexico double or triple] ; 7-P usually double, rare-
ly single (1-3); 4-M usually double (1-3). Abdomen: Hairs 10,1 1-I subequal in length;
3-III usually double (1-2); 5-VII more than half length of 3-VII, often subequal to 3-
VIL. Segment VIII: Comb scales 56 to more than 100 (highest number in only Mexi-
can specimen counted); length of free portion of midapical scale 0.027-0.035 mm
(0.025-0.037 mm). Siphon: Length 0.72-0.90 mm (Mexican specimens not meas-
ured); L/S 2.04-2.22 (1.96-2.33); P/L 0.53-0.56 (0.51-0.58); H/L 0.59-0.63 (0.57-
0.64). Anal Segment: Ventral brush usually with 14 hairs (12-15, rarely 16); 4a-X
usually 6-branched (5-6, rarely 4 or 7); dorsal gill at least subequal in length to
saddle.
SYSTEMATICS. Aedes homoeopus differs from impostor, the only other mem-
ber of the subgroup, in the structure of the genitalia. In homoeopus the median
sternomesal area usually shows a well-developed convexity, sclerite and tuft and the
filament of the prosophallus is usually strongly angulate; in impostor the median
sternomesal specializations are poorly differentiated and the filament is apparent-
ly only weakly angulate. The larvae of homoeopus and impostor are similar but
show the following differences. In homoeopus hairs 10,11-I are subequal in length
and 4a-X is usually 6-branched, rarely 7-branched; in impostor 10-I is at least 1.3
the length of 11-I and 4a-X is 8-branched (1 specimen).
The male of homoeopus shows some marked variation in diagnostic characters.
The dorsocentral area at one extreme is silvered for nearly its entire length and at
the other extreme the dorsocentral area is entirely devoid of silver scales (GUA
48-11); there are intermediate conditions in which the dorsocentral area is anter-
iorly silvered to various extents caudad. When it is not extensively silvered the
acrostichal and usually the posterior dorsocentral stripes persist as prominent mark-
ings. The mesonotum of the GUA 48-11 male is strikingly similar to that of het-
eropus (figs. 31,42). The median sternomesal convexity and sclerite occasionally
show marked reduction from the strongly developed condition typical of homoe-
opus and in 2 specimens from Guatemala, the hook of the claspette filament, typi-
cally strongly angulate, is apparently only weakly angulate. The median sterno-
mesal structures of the latter 2 specimens are also reduced and only the presence
of the median sternomesal tuft serves to distinguish these from impostor on the
basis of the genitalia.
DISTRIBUTION (fig. 4). State of Veracruz, Mexico, at elevation of 3000 ft; Gua-
temala south to Costa Rica at elevations of 3100 to possibly 5000 ft. Material exam-
ined: 282 specimens; 82 6, 89 2, 52 pupae, 59 larvae; 49 individual rearings (36
larval, 10 pupal, 3 incomplete).
COSTA RICA. Alajuela: Alajuela [elev. 3100 ft], 26 May 1921, A. Alfaro, 1 2?;June 1921, bam-
boo, A. Alfaro, 1 2; 1 July 1921, bamboo, A. Alfaro, 1 9; 4 July 1921. bamboo, A. Alfaro, |g;
30 July 1921, A. Alfaro, 2 9; Aug 2k bamboo, A. Alfaro, 1 d, type series in part (1545, USNM
25253).9 Oct 1921, A. Alfaro, 3 6, ye FiolZ Oct 1921; A. hate 2 6% Oct 192) Av Alfaro.
Lad,.2 9; Apr 1922, A. Alfaro, 2 ?, type series in part (USNM 25253) [USNM]. Alajuela [elev.
3100 fi 123 km S, 13 Nov 1962, Gaies stumps, C. Hogue, W. Powder (CR 18), 2 lpd (18-103,
105), 2 Ip? (18-106,107), 1 L (18-2) [UCLA]. San Jose: Guadalupe [elev. 4000 ft], 27 June
1922, A. Alfaro, 1 ? [USNM]. San Jose, Parque Bolivar, elev. 3800 ft, 8 June 1963, small treehole,
height 7 ft, C. Hogue (CR 88), 6 lp? (88-102,104,105,107,109,110), 2 pd (88-202,203), 2 Ip? (88-
66 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
201), 4d, 4 9, 24 L (88-1) [UCLA] . San Jose, University of Costa Rica, elev. 3800 ft, 8 Nov 1962,
large treehole, J. Belkin, C. Hogue, W. Powder (CR 5), 1 Ipd (5-111), 7 Ip? (5-101-106,110), 1 Ip
(5-107), 1 L (5-1) — treehole, 1 Ip? (CR 6-101); 13 Nov 1962, treehole, J. Belkin, C. Hogue, W.
Powder, 1 lp? (CR 15-101); 16 Nov 1962, treehole (same as CR 5), C. Hogue, W. Powder, 1 Ipd
(CR 25-101) — large treehole (CR 44), 1 Ipd (44-102-103), 1 Ip? (44-101), 1 IP (44-104); 14 June
1964, large treehole, height 6 ft, C. Hogue (CR 175), 7 lp? (175-101,102,104-107,110), 5 p?
(175-201-205), 1 6, 1 9, 2 p, 3 L (175-3) [UCLA]. Province not specified: Miravalles (? Guana-
casta: Hacienda Miravalles, elev. 1700-3300 ft), 20 July 1922, A. Alfaro, 1 9 [USNM].
EL SALVADOR. Department not specified: El Boqueron, 2 6 (716), 1 6 (717), 6 6, 8°
[USNM].
GUATEMALA. Chimaltenango: Aldea “Pachup”, nearest town Yepocapa [elev. 4850 ft], 28
Sept 1950, treehole, R. Rodriquez, P. Coc and J. Giron, 24 6, 1 9 (GUA 149-1) [UCLA] . Finca
Santa Izabel, nearest town Yepocapa [elev. 4850 ft], 3 Aug 1950, treehole, V. and M. Xinic, 2,
3 9 (GUA 143-1) [UCLA]. Rio Queleya, nearest town Yepocapa [elev. 4850 ft], 22 Sept 1950,
un pocito de agua estancada, J. and G. Giron, 1 ¢ (GUA 147-2) [UCLA]. Locality not specified,
nearest town Yepocapa [elev. 4850 ft], 20 July 1950, treehole, P. Garcia, M. Xinic, 14. 6,7 %
(GUA 140-2); 29 July 1950, treehole, H. Dalmat, 4 6 (GUA 141-2); 3 Aug 1950, treehole, A.
Castellanos, P. Garcia, 5 9? (GUA 142-1) [UCLA]. Guatemala: Amatitlan, elev. ca 3900 ft, 16 July
1964, small treehole, height 5 ft, P. Cowsill (GUA 48), 3 Ipd (48-10,11,13), 1 IP (48-12), 1 2, 1 p,
1 L (48-1) [UCLA]. Guatemala, Villa de Guadelupe cemetery, elev. ca 5000 ft, 3 Sept 1964, small
cement trough, W. Almengor, P. Cowsill (GUA 122), 1 p? (122-101), 1 2? (122-100) [UCLA].
MEXICO. Veracruz: Cordoba [elev. 3000 ft], 7 Mar 1908, treehole, F. Knab (429), 2 Ip?
(429.11,.31), 2 6, type series in part (429.3, USNM 25253; 429.9, without USNM number), 2 6
(429.14), 5 6(429p), 1 6 genitalia (1543, adult number unknown), 6 ? (429 .4,.14,.20,.23,.59,.62),
2 2 (429p) [USNM] ;7 Aug 1905, biting-landing, 1500-1600 hrs, R. Schick, D. Schroeder, A. Lau,
1 9 (MEX 278-1) [UCLA].
15. Aedes (Finlaya) impostor Schick, n.sp.
Figs. 4,33,34
TYPE. Holotype 6 (GUA 33-30) with associated pupal and larval skins and genitalia slide, Finca
Trece Aguas, elev. 2800 ft, Alta Vera Paz, Guatemala, 7 July 1964, cut or broken bamboo, T. and
J. Zavortink [USNM] . Paratypes: 5 L (33-3), same data as holotype [UCLA].
MALE. Head: Vertex with all decumbent scales silver, narrow curved; palpus
about 2 labellum lengths shorter than proboscis, segment 4 with hairs of ventrolat-
eral row closely spaced. Thorax (fig. 33): Mesonotal disc transversely silvered for
more than half length. Legs: Tarsus 1-IJ with median dark band about 0.5. Wing:
Vein Cu with long line of silver scales on stem.
MALE GENITALIA (fig. 33). Sidepiece: Length 0.42 mm; median sternomesal
area with convexity and sclerite weakly developed, tuft absent. Prosophallus: Length
0.14:mm; mesal lobe with lateral portion inclined at about 30° from horizontal;
filament ratio 0.73, hook not strongly angulate. Aedeagus: Length 0.14 mm.
PUPA (fig. 33). Cephalothorax: Hair 9-C double. Abdomen: Hair 1-I with pri-
mary branches predominantly single; 2-II laterad of 3-II. Paddle: Ventral midrib
weakly pigmented.
LARVA (fig. 34). Head: Hair 5-C usually single (1-2); 6-C single; 7-C usually triple
or 4-branched (3-5). Thorax: Hair 1-P usually triple (2-4); 4-P usually double or trip-
le (2-4); 5-P usually double (2-3); 7-P usually triple (2-3); 4-M double. Abdomen:
Hair 10-I 1.3 or more length of 11-I; 3-III double; 5-VII usually less than half length
Schick: Terrens Group of Aedes (Finlaya) 67
of 3-VII, at most slightly longer than half length. Segment VIII: Comb scales 50 to
more than 100; length of free portion of midapical comb scale 0.027-0.030 mm. Si-
phon: Length 0.91-1.00 mm; L/S 2.37-2.44; P/L 0.56-0.59; H/L 0.63-0.66. Anal
Segment: Ventral brush with 11-13 hairs; 4a-X 8-branched; dorsal gill absent in all
specimens.
SYSTEMATICS. Aedes impostor is very similar to homoeopus. Their separation
is discussed under the latter species.
DISTRIBUTION (fig. 4). Department of Alta Vera Paz, Guatemala. Material ex-
amined: 8 specimens; 1 6, 1 pupa, 6 larvae; | individual rearing (larval).
GUATEMALA. Alta Vera Paz: Finca Trece Aguas, elev. 2800 ft, 7 July 1964, cut or broken
bamboo, T. and J. Zavortink (GUA 33, type series), 1 lpd (33-30), 5 L (33-3) [USNM; UCLA].
Heteropus Subgroup
ADULTS. Head: Vertex of both sexes with decumbent scales along longitudinal
midline narrow; occiput of females with erect scales variable, from all pale to all
dark, of males, all pale; proboscis of females shorter or longer than femur I; palpal
segments 3 and 4 of males with ventrolateral hairs variously developed. Thorax: Mes-
onotal disc of both sexes not transversely silvered; complete acrostichal or posterior
dorsocentral lines present or absent in females; acrostichal seta present; fossal mac-
ula of females not coextensive with fossa, reduced mesally, of males coextensive
or not coextensive; supraalar macula not truncate posteriorly, in females usually
not joined to fossal macula; ppn silver scaled; ssp scales present or absent; pra
hairs of females pale; upper stp and mep scale patches not contiguous. Legs: Mid-
and hindlegs not shaggy; mid- and hindtarsi with conspicuous silver bands; tarsus
1-[ with or without narrow apical band; femur II with knee spot broad, the scales
extending basad of anterior subapical setae; tarsus 1-II with median dark band in-
complete or complete and moderately broad; tarsus 2-II usually with complete dark
apical band; tarsus 5-II without silver scales; femur II] with basal dark band incom-
plete or complete and usually narrow; tarsus 5-III without silver scales. Wing: Vein
C of female usually with small patch of silver scales or line not reaching 0.5 to cross-
vein h, of males usually a small patch, sometimes an interrupted line reaching h;
veins R and Cu of males without silver scales.
MALE GENITALIA. Sidepiece: Basal tergomesal area with or without dense patch
of long setae, median sternomesal area with or without strongly developed sclerite,
with or without tuft; specialized subapical sternal seta absent. Prosophallus: Mesal
lobe with lateral portion inclined from less than 15° to about 45° from horizontal;
stem not bowed; filament with hook strongly or not strongly angulate.
PUPAE. Cephalothorax: Without pale inverted V-shaped marking; hair 5-C less
than twice length of 4-C; 9-C usually single. Abdomen: Hair 1-I with primary branch-
es predominantly single to multiple; 2-IJ usually mesad of 3-II for 0.3 or less the dis-
tance from 1-II to 3-II; 3-III single. Paddle: Pigmented; apex usually produced, often
strongly; hair 1-P shorter than paddle.
LARVAE. Head: Hairs 5,6-C single to multiple: 7-C more than half length of 6-C;
11-C longer than mental plate; 14-C longer than mental plate, single; bmh single or
branched. Antenna: Hair 1-A single. Thorax: Hair 11-P greater or less than half
length of 14-P; 14-P single; 3-M single; 4-M usually double; 8-T shorter, but more
than half length of, or longer than metathoracic pleural tubercle. Abdomen: Hair 5-I
at least subequal in length to 4-I; 2-II well mesad of 4-II; 3-III single or double;
68 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
10-III single; 3-VI single; 4-VII usually single; 5-VII caudad of 4-VII, shorter or
longer than half length of 3-VII; 10,12-VII single. Segment VIII. Midapical comb
scale with free portion longer or shorter than sessile portion; 2-VIII single. Anal
Segment: Saddle extending around segment for moderate distance, ventral margin
without deep slit. |
DISCUSSION. The Heteropus Subgroup is a rather large and diverse group but
the species show no apparent fundamental differences that would justify the recog-
nition of further subgroups. The subgroup is characterized in the adults by the
combination of the absence of transverse silvering on the mesonotum of the males,
the presence of acrostichal setae, the disjunct fossal and supraalar maculae of the
females, the pale pra hairs, the incomplete or narrow dark basal band of femur III
and the absence of extensive silver scaling at the base of the wing veins. The larva
of some species shows nonmultiple hairs 5,6-C and cannot be readily distinguished
from those of the Metoecopus, Homoeopus and Podographicus Subgroups; in the
remaining species the larvae are distinguished by the combination of characters giv-
en in the key. The male genitalia and pupae do not show distinctive subgroup
characters.
The subgroup occurs at moderately high to high elevations from Mexico to Costa
Rica.
Three phyletic lines appear to be represented, one by amabilis, gabriel, idanus
and sumidero, a second by vargasi and a third by heteropus. The species of the am-
abilis line seemingly are allied by the mesonotal markings of the females, complete
acrostichal and/or posterior dorsocentral lines tending to be developed and the
fossal maculae being strongly concave mesally. 7
16. Aedes (Finlaya) amabilis Schick, n.sp.
Figs: 5,37
TYPE: Holotype 2 (MEX 381-3), Cueva del Nacimiento del Agua (cave of origin of Rio
Atoyac), nearest town Penuela (elev. 3000 ft), Veracruz, Mexico, 13 July 1965, biting-landing,
C.L. Hogue [USNM] . Paratypes: 3 ? (381-3), same data as holotype [BM; UCLA].
FEMALE. Head: Vertex with decumbent scales laterad of median longitudinal
line generally dark, narrow curved; occiput with erect scales dark; proboscis longer
than femur I. Thorax (fig. 37): Complete acrostichal and posterior dorsocentral
lines present; most posterior acrostichal seta at about 0.5 from anterior end or in
posterior half; fossal macula with mesal margin concave; supraalar macula not join-
ed to fossal macula: prescutellar space and midlobe of scutellum with silver scales;
ssp scales present. Legs: Femur I with narrow knee spot; with poorly developed
posterior patch of silver scales in basal half comprising scattered scales; posterior
patch of femur II either similarly developed or with fewer scales or absent; tarsus
1-II with median dark band complete, about 0.5; femur III with basal dark band
0.07-0.11 (incomplete in 1 specimen), subapical dark band 0.36-0.42; tarsus 1-III
with basal silver band 0.13-0.16, apical silver band 0.14-0.21. Wing: Vein C with
small basal patch of silver scales or without silver scales.
MALE, PUPA, LARVA. Unknown.
SYSTEMATICS. Aedes amabilis is distinguished from the other members of the
subgroup by the presence of both complete acrostichal and posterior dorsocentral
Schick: Terrens Group of Aedes (Finlaya) 69
lines and by the presence of a narrow knee spot on femur I.
DISTRIBUTION (fig. 5). State of Veracruz, Mexico, at elevation of 3000 ft. Ma-
terial examined: 6 2; no individual rearings.
MEXICO. Veracruz: Cueva del Nacimiento del Agua (cave of origin of Rio Atoyac), nearest
town Penuela (elev. 3000 ft), 13 July 1965, biting-landing, C. Hogue, 4 9 (MEX 381-3, type ser-
ies) [USNM; BM; UCLA]. Presidio, 0.5 mi S, elev. 2300-3300 ft, 14 July 1965, biting-landing,
1500 hrs, C. Hogue, 2 9? (MEX 386-3) [UCLA].
17. Aedes (Finlaya) gabriel Schick, n.sp.
Figs: 5.35.36
TYPE: Holotype 36 (MEX 350-113) with associated pupal skin, Gabriel Mariaca, 0.5 mi E (2.7
mi W Tepotzlan), elev. 5200 ft, Morelos, Mexico, 7 Sept 1965, large treehole, D. Schroeder
[USNM]. Allotype ? (350-20) with associated pupal and larval skins, same data as holotype
[USNM] . Paratypes: 1 Ipé (350-23), 12 lpd (350-10,18-22,26,28 29), 10 pd (350-92,95,100,101,
103,104,108-110,114), 7 p? (350-90,91,94,102,105,107,111), 6 36, 4 9, 35 P, 9 p, 50 L (350-1),
same data as holotype [BM; UCLA].
? Aedes (Gualteria) terrens in part of Vargas and Downs (1950:171).
FEMALE. Head: Vertex with decumbent scales laterad of median longitudinal
line silver or dark, narrow curved; occiput with erect scales all pale or all dark or
with a mixture of both; proboscis shorter than or subequal in length to femur I.
Thorax (fig. 35): Complete acrostichal line present: complete posterior dorsocen-
tral line absent; most posterior acrostichal seta usually at about 0.25 from anterior
end, but sometimes in posterior half; fossal macula with mesal margin concave:
supraalar macula not joined to fossal macula; prescutellar space and midlobe of
scutellum with silver scales; ssp scales present. Legs: Femur I without knee spot;
femora I and II with well-developed posterior patch of silver scales in basal half,
about equal in size; tarsus 1-II with median dark band complete, about 0.33-0.5;
femur III with basal dark band 0.05-0.08, subapical dark band 0.35-0.49; tarsus
1-I]I with basal silver band 0.07-0.11, apical silver band 0.14-0.21 (0.14-0.25). Wing:
Vein C with small basal patch of silver scales; vein R without silver scales.
MALE. Head: Vertex with all decumbent scales silver, narrow curved; palpus sub-
equal in length to proboscis or 0.5 labellum length longer; segment 3 with 3 or fewer
rather short apical ventrolateral hairs; segment 4 with hairs of ventrolateral row
short, sparse. Thorax (fig. 35): Mesonotal disc without strip of silver scales bordering
fossa; complete acrostichal line usually present, middle portion sometimes weakly
developed; complete posterior dorsocentral line absent; fossal macula not reaching
mesal margin of fossa. Legs: Tarsus 1-II with median dark band as in female. Wing:
Vein C with small basal patch of silver scales.
MALE GENITALIA (fig. 35). Sidepiece: Length 0.43-0.45 mm; basal tergomesal
area without dense patch of long setae; median sternomesal area with convexity usu-
ally weakly, sometimes moderately, developed, sclerite well developed, tuft present.
Prosophallus: Length 0.15 mm; mesal lobe with lateral portion inclined from about
30° to 45° from horizontal; filament ratio 0.35-0.40; hook of filament not strongly
angulate. Aedeagus: Length 0.15-0.16 mm.
PUPA (fig. 35). Cephalothorax: Hair 9-C single. Abdomen: Hair 1-I with primary
branches predominantly single or single-double; 2-II laterad of 3-II to mesad for
70 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
more than 0.3 the distance from 1-II to 3-II, usually mesad for 0.3 or less. Paddle:
Apex not or weakly to moderately produced; ventral midrib usually weakly pigment-
ed, when infrequently moderately pigmented not more strongly at apex.
LARVA (fig. 36). Head: Hairs 5,6-C single; 7-C usually double to 4-branched (2-
5); bmh single. Thorax: Hair 1-P usually double, rarely triple; 4-P single or double; 5-
P usually double, rarely triple; 7-P usually double, rarely triple; 11-P less than half
length of 14-P; 4-M double; 8-T usually shorter than metathoracic pleural tubercle
(subequal in length in MEX 352). Abdomen: Hair 2-I usually single (1-2); 3-III usu-
ally single (1-2). Segment VIII: Comb scales 80 to more than 100, usually the latter;
free portion of apical scale shorter or longer than sessile portion, length 0.015-0.030
mm (0.024-0.033 mm). Siphon: Length 0.70-0.80 mm; L/S 1.89-2.00 (1.82-2.08);
P/L 0.57-0.61 (0.56-0.63); H/L 0.62-0.67 (0.61-0.68). Anal Segment: Ventral brush
usually with 12 hairs (11-13); 4a-X usually 5-branched (4-6).
SYSTEMATICS. Aedes gabriel is distinguished from the other members of the
subgroup in the adults by a combination of the presence of a complete acrostichal
line and the absence of a complete posterior dorsocentral line; in the male gen-
italia by a combination of the absence of a dense patch of long basal tergomesal
setae and the presence of a well-developed median sternomesal sclerite and tuft on
the sidepiece; in the larva by a combination of the single hairs 5,6-C, the usually
single 2-I and the ventral brush of usually 12 hairs.
The only larva of lot MEX 352 (352-12), differs markedly from the other gabriel
larvae in characters that would otherwise be of some use in separating this species
from the others in which hairs 5,6-C are not multiple and 4-VII is single. Typically,
in gabriel, hair 8-T is shorter than the metathoracic pleural tubercle, 10-I is much
longer than 11-I, hairs 1,13-III-VI are single and long and 5-VII is shorter than 3-
VI. In MEX 352-13, however, hair 8-T is subequal in length to the tubercle, 10,11-I
are subequal in length, 1,13-III-VI are multiple and relatively short and 5-VII is
longer than 3-VII. The pupa of this rearing unfortunately died so that the adult is
not available for confirmation of the identification.
DISTRIBUTION (fig. 5). State of Morelos, Mexico, at elevation of 5200 ft. Mater-
ial examined: 234 specimens; 30 6, 30 9, 87 pupae, 87 larvae; 50 individual rearings
(21 larval, 26 pupal, 3 incomplete).
MEXICO. Morelos: Gabriel Mariaca, 0.5 mi E (2.7 mi W Tepotzlan), elev. 5200 ft, 7 Sept 1965,
volcanic rockhole, D. Schroeder, 1 pd (MEX 346-102) — small treehole (MEX 349), 4 lp? (349-
13-15,17), 3 pd (349-102-104), 1 IP (349-12), 1 6, 2 L (349-1) [UCLA] — large treehole (MEX
350, type series), 1 Ipd (350-23), 13 Ip? (350-10,18-22,26,28,29), 11 pd (350-92,95,100,101,103,
104,108-110,113,114), 7 p? (350-90,91,94,102,105,107,111), 6 6, 4 2, 35 P, 9 p, 50 L (350-1)
[USNM; BM; UCLA] — small treehole (MEX 352), 1 pd (352-101), 1 p? (352-100), 1 IP (352-13),
1 L (352-2) — small volcanic rockhole (MEX 355), 1 Ipd (355-10), 2 pé (355-100,102), 2 IP (355-
Lb,12), 1 L 55-1) [UCLA].
18. Aedes (Finlaya) idanus Schick, n.sp.
Figs. 537,38
TYPE: Holotype 6 (MEX 420-10), with associated larval and pupal skins and genitalia slide,
Viejo de Agua de Obispo, nearest town Chilpancingo (elev. 2220 ft), Guerrero, Mexico, 8 Aug
1966, treehole, D. Schroeder [USNM]. Allotype 9 (MEX 413-1), Agua de Obispo, elev. 2900 ft,
8 Aug 1966, biting, 0800-0830 hrs, D. Schroeder [USNM]. Paratypes: 3 L (420-1), same data as
holotype; 1 ? (413-1), same data as allotype [UCLA].
Schick: Terrens Group of Aedes (Finlaya) 71
FEMALE. Head: Vertex with decumbent scales of area just adjacent to median
longitudinal line dark, narrow curved; scales laterad of these dark, broad; occiput
with erect scales pale or dark; proboscis longer than femur I. Thorax: Complete
acrostichal line absent although line of scattered silver scales present in specimens
from Taxco; complete posterior dorsocentral line present; most posterior acrosti-
chal seta about 0.4 from anterior end; fossal macula with mesal margin essentially
straight; supraalar macula not joined to fossal macula; prescutellar space and mid-
lobe of scutellum with silver scales; ssp scales present. Legs: Femur I without knee
spot; femora I and II with well-developed posterior patch of silver scales in basal
half, that of femur II larger; tarsus 1-II with median dark band complete, about 0.4-
0.6; femur III with basal dark band incomplete, subapical dark band 0.31-0.34; tar-
sus 1-III with basal silver band 0.15, apical silver band 0.19. Wing: Vein C with small
basal patch of silver scales; vein R without silver scales.
MALE. Head: Vertex with decumbent scales laterad of median longitudinal line
dark, broad; palpus about 2 labellum lengths shorter than proboscis; segment 3 with
3 short apical ventrolateral hairs; segment 4 with hairs of ventrolateral row short and
sparse. Thorax (mesonotal disc largely rubbed): Fossal macula not reaching mesal
margin of fossa. Legs (in poor condition, measurement possibly inaccurate): Tarsus
1-II with median dark band complete, about 0.6. Wing: Vein C with small basal
patch of silver scales.
MALE GENITALIA (fig. 37). Sidepiece: Length 0.38 mm; basal tergomesal area
with dense patch of long setae; median sternomesal area with convexity moderately
developed, sclerite well developed, tuft present. Prosophallus: Length 0.11 mm; mes-
al lobe with lateral portion inclined at about 30° from horizontal; stems essentially
parallel; filament ratio 0.50-0.55; hook of filament strongly angulate. Aedeagus:
Length 0.14 mm.
PUPA (fig. 37). Cephalothorax: Hair 9-C single. Abdomen: Hair 1-I with primary
branches predominantly single; 2-II mesad of 3-II for less than 0.3 the distance from
1-II to 3-IIl. Paddle: Apex moderately produced; ventral midrib moderately pig-
mented, not more strongly at apex.
LARVA (fig. 38). Head: Hair 5-C 4-6 branched; 6-C single or double; 7-C 5-7
branched; bmh single. Thorax: Hair 1-P triple; 4-P double or triple; 5-P double; 7-P
double; 11-P less than half length of 14-P; 4-M double; 8-T shorter than metathoracic
pleural tubercle. Abdomen: Hair 2-I single to triple; 3-III single or double. Segment
VII: Comb scales about 42; length of free portion of midapical scale 0.033 mm
(length relative to sessile portion not known). Siphon: Exact length not determined
because of poor orientation on slide, at least 0.91 mm. Anal Segment: Ventral brush
with 12 hairs; 4a-X 6-branched.
SYSTEMATICS. Aedes idanus is distinguished from the other members of the
subgroup in the female by a combination of the long proboscis, the absence of a
complete acrostichal line and the presence of a complete posterior dorsocentral
line; in the male genitalia by the combination of a patch of long basal tergomesal
setae and a well-developed median sternomesal sclerite and tuft of the sidepiece;
and in the larva by the multiple hair 5-C and the short 11-P.
DISTRIBUTION (fig. 5). State of Guerrero, Mexico, at elevations of 2220 or
3000-5800 ft. Material examined: 10 specimens; 1 6, 4 9, 1 pupa, 4 larvae; 1 andivid-
ual rearing (larval).
MEXICO. Guerrero: Agua de Obispo (21 mi S Chilpancingo), elev. 2900 ft [possibly 3000-4000
ft], 8 Aug 1966, biting, 0800-0830 hrs, D. Schroeder, 2 9 (MEX 413-1, type series in part)
[USNM; UCLA]. Rancho Viejo de Agua de Obispo (24 mi S Chilpancingo), elev. 2200 ft [pos-
72 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
sibly 3000-4000 ft], 8 Aug 1966, large treehole in dead tree, height 10 ft, D. Schroeder (MEX
420, type series in part), 1 lp? (420-10), 3 L (420-1) [USNM; UCLA] . Tazco (Taxco) [elev. 5800
ft], Aug 1955, N. Krauss, 2 9 [USNM].
19. Aedes (Finlaya) sumidero Schick, n.sp.
Figs. 5,39
TYPE: Holotype 6 (MF 8-2), Sumidero (24 km N Tuxtla Gutierrez), elev. 3400 ft, Chiapas,
Mexico, 16 Aug 1964, in shelter flying at night, E. Fisher [USNM]. Allotype ° (MF 8-2), same
data as holotype [USNM]. Paratypes: 12 6,4 9 (MF 8-2), same data as holotype [BM; UCLA].
FEMALE. Head: Vertex with decumbent scales of area just adjacent to median
longitudinal line variable, usually dark, narrow curved; scales laterad of these sil-
ver, broad; occiput with erect scales all pale or with mixture of pale and dark scales;
proboscis shorter than femur I. Thorax (fig. 39): Complete acrostichal line absent;
complete posterior dorsocentral line absent or weakly developed; most posterior
acrostichal seta at about 0.5 from anterior end; fossal macula with mesal margin
concave; supraalar macula usually not joined to fossal macula (narrowly joined in
1 specimen); prescutellar space and midlobe of scutellum with silver scales; ssp
scales present. Legs: Femur I without knee spot; femora I and II with well-develop-
ed posterior patch of silver scales in basal half; subequal in size; tarsus 1-II with
median dark band complete, usually about 0.5, sometimes about 0.4; femur III
with basal dark band usually incomplete, when complete 0.05 or less, subapical
dark band 0.33-0.36 (0.30-0.37); tarsus 1-III with basal silver band 0.11-0.16 (0.10-
0.19), apical silver band 0.17-0.22 (0.17-0.23). Wing: Vein C with small basal patch
of silver scales; vein R without silver scales.
MALE. Head: Vertex with all decumbent scales silver, narrow curved; palpus
usually about 0.5 labellum length shorter than proboscis, sometimes 1.5 less; seg-
ment 3 with several short to moderately long apical ventrolateral hairs shorter
than segments 4 and 5 combined; segment 4 with hairs of ventrolateral row short to
moderately long, somewhat sparse. Thorax (fig. 39): Mesonotal disc with strip of sil-
ver scales bordering fossal macula much broader posteriorly than anteriorly; com-
plete acrostichal line usually absent, sometimes weakly developed; complete poster-
ior dorsocentral line present; fossal macula coextensive with fossa. Legs: Tarsus |-II
with median dark band complete, about 0.4. Wing: Vein C with basal silver scales
variably developed, often forming incomplete line reaching crossvein h.
MALE GENITALIA (fig. 39). Sidepiece: Length 0.41-0.46 mm; basal tergomesal
area without dense patch of long setae; median sternomesal area with convexity and
sclerite weakly developed, tuft present but not prominently differentiated. Proso-
Phallus: Length 0.14-0.15 mm; mesal lobe with lateral portion usually inclined at
15° or less than 15° from horizontal (unilaterally between 15° and 30° in 1 speci-
men); stems essentially parallel; filament ratio 0.50-0.60; hook of filament not
strongly angulate. Aedeagus: Length 0.14-0.15 mm.
PUPA, LARVA. Unknown. |
SYSTEMATICS. Aedes sumidero is distinguished from the other members of the
subgroup in the female by a combination of the short proboscis and the absence
of a complete mesonotal line; and in the male by the form of the anterolateral sil-
ver strip of the mesonotal disc, which is unique for the Terrens Group. The male
Schick: Terrens Group of Aedes (Finlaya) 73
genitalia serve to separate sumidero from gabriel, idanus and heteropus but not from
vargasi; the sidepiece lacks a basal tergomesal tuft of long setae and prominent spec-
ialization of the median sternomesal area and the lateral portion of the mesal lobe
of the prosophallus is only slightly inclined.
DISTRIBUTION (fig. 5). State of Chiapas, Mexico, at elevation of 3400 ft. Mater-
ial examined: 45 specimens; 18 6, 27 9; no individual rearings.
MEXICO. Chiapas: Sumidero (24 km N Tuxtla Gutierrez), elev. 3400 ft, 23 July 1963, biting-
landing, 0900 hrs, E. Fisher, 13 6, 5 9? (MF 8-2, type series) [USNM; BM; UCLA]; 15 Aug 1964,
“in shelter flying at night”, E. Fisher, 3 2? (MEX 115-2); 17 Aug 1964, biting-landing, 1100 hrs,
E. Fisher, D. Verity (MEX 120-128), 1 ° (120-2), 4 9 (121-3), 6 2 (122-1; 123-1; 124-1; 125-1;
126-1; 127-1), 5 6) 7-9.(128-3) [UCLA].
20. Aedes (Finlaya) vargasi Schick, n.sp.
Figs. 5,40,41
TYPE: Holotype 3 (MEX 247-10) with associated pupal and larval skins and genitalia slide,
Cordoba, 5-15 mi S, elev. 3000-3500 ft, Veracruz, Mexico, 30 July 1965, treehole, D. Schroeder
[USNM]. Allotype ? (MEX 246-10) with associated pupal and larval skins, same data as holotype
[USNM]. Paratypes: 1 P, 22 L (247-1), same data as holotype [BM; UCLA]. This species is dedi-
cated to Luis Vargas of the Instituto de Salubridad y Enfermedades Tropicales, Mexico.
FEMALE. Head: Vertex with decumbent scales laterad of median longitudinal
line silver or dark, narrow curved; occiput with erect scales pale. Thorax (fig. 40):
Complete acrostichal or posterior dorsocentral lines absent; most posterior acrost-
ichal seta usually at about 0.25 from anterior end (in MEX 228-10 at poster-
ior end); fossal macula with mesal margin essentially straight or slightly concave;
supraalar macula not joined to fossal macula; prescutellar space and midlobe of
scutellum with silver scales; ssp scales usually absent (MEX 226-10 with 2 scales).
Legs: Femur I without knee spot; femora I and II with well-developed posterior
patch of silver scales in basal half, about equal in size; tarsus 1-II with median dark
band usually complete, about 0.33 (incomplete in 1 specimen of MEX 18); femur
III with basal dark band 0.07-0.13, subapical dark band 0.30-0.35; tarsus 1-III with
basal silver band usually present, 0.04-0.06 (absent in MEX 346-10), apical silver
band 0.21-0.31. Wing: Vein C with basal silver scales forming small patch or line not
reaching 0.5 to crossvein h; vein R with a few basal silver scales.
MALE. Head: Vertex with all decumbent scales silver, narrow curved; palpus a-
bout 1 labellum length shorter than proboscis; segment 3 with several long apical
ventrolateral hairs forming tuft nearly as long as segments 4 and 5 combined; seg-
ment 4 with hairs of ventrolateral row increasing in length basad, sparse. Thorax (fig.
40): Mesonotal disc without strip of silver scales bordering fossal macula; complete
acrostichal or posterior dorsocentral lines absent; most posterior acrostichal seta
about 0.25 from anterior end; fossal macula coextensive with fossa. Legs: Tarsus 1-
If with median dark band virtually obliterated by silver scales. Wing: Vein C with
basal silver scales forming line reaching 0.5 to crossvein h.
MALE GENITALIA (fig. 40). Sidepiece: Length 0.30 mm; basal tergomesal area
without dense patch of setae; median sternomesal area with convexity absent, scler-
ite weakly developed, tuft absent. Prosophallus: Length 0.11 mm; mesal lobe with
lateral portion inclined at less than 15° from horizontal; stems slightly divergent; fil-
ament ratio 0.50-0.55; hook of filament not strongly angulate. Aedeagus: Length
O12 mm.
74 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
PUPA (fig. 40). Cephalothorax: Hair 9-C usually single, infrequently double. Ab-
domen: Hair 1-I with primary branches predominantly double-triple to predomin- —
antly multiple, infrequently predominantly single (but with long barbs); 2-II mesad
of 3-II for usually 0.3 or less the distance from 1-II to 3-II, sometimes more than 0.3.
Paddle: Apex usually strongly produced; ventral midrib moderately pigmented for
most length, weakly at apex.
LARVA (fig. 41). Head: Hair 5-C usually single (1-3); 6-C single; 7-C double to 5-
branched; bmh single. Thorax: Hairs 1,4-P usually double or triple (2-4); 5-P single
or double; 7-P usually double, infrequently triple; 11-P less than half length of 14-P;
8-T shorter than or subequal in length to metathoracic pleural tubercle. Abdomen:
Hair 2-I usually triple (2-3); 3-III usually double (1-2). Segment VIIT: Comb scales
40 to more than 100 [in Tamazunchale 40-60; in Cordoba 50 to more than 100];
free portion of midapical scale usually longer than sessile portion, length 0.027-
0.033 mm (0.022-0.036 mm). Siphon: Length 0.80-0.97 mm; L/S 2.13-2.56 [in Cor-
doba 2.13-2.44; in Tamazunchale 2.27-2.56]; P/L 0.55-0.59 (0.54-0.60); H/S 0.61-
0.65 (0.60-0.67). Anal Segment: Ventral brush usually with 14 hairs (13-15); 4a-X
usually 6,7-branched (5-8).
SYSTEMATICS. Aedes vargasi is distinguished from the other members of the
subgroup in the adults by the absence of complete acrostichal and posterior dorso-
central lines in both sexes, the absence of a scale patch on the ssp and the usually
narrower median dark band of tarsus 1-II; in the larva by a combination of the
single hairs 5,6-C, the usually branched 2-I and the ventral brush of usually 14
hairs. The male genitalia are similar to those of sumidero.
DISTRIBUTION (fig. 5). Atlantic drainage of Sierra Madre Oriental, Mexico, at
elevations of 400 to about 3000 ft. Material examined: 63 specimens; 1 6, 10 9,9
pupae, 43 larvae; 9 individual rearings (6 larval, 2 pupal, 1 incomplete).
MEXICO. San Luis Potosi: Saketepan (near Tamazunchale), elev. 400 ft, 20 July 1965, small
treehole, height 6 ft, R. Schick, D. Schroeder (MEX 213), 1 p? (213-100), 9 L (213-3) — biting-
landing, 1100 hrs, 1 ? (MEX 220-7) — small treehole, height 10 ft, 11 L (MEX 227-1) — small tree-
hole, height 3 ft, 1 lp? (MEX 228-10) — small treehole, height 4 ft (MEX 229), 2 lp? (229-10,11),
1 p? (229-100) [UCLA] . Veracruz: Cordoba, 1.5 mi E, elev. 3000 ft, 12 July 1964, biting-landing,
1900 hrs, at dusk, E. Fisher, D. Verity, 2 9 (MEX 18-1); 22 July 1964, large treehole, height 5 ft,
E. Fisher, D. Verity, 1 lp? (MEX 71-5) [UCLA]. Cordoba, 5-15 mi S, elev. 3000-3500 ft, 30 July
1965, small treehole, height 5 ft, D. Schroeder (MEX 245), 1 IP (245-10), 1 L (245-1) [UCLA] —
small treehole (MEX 246), 1 lp? (246-10), 3 L (246-1) [USNM; UCLA] — small treehole, height 5
ft (MEX 247, type series), 1 lpd (247-10), 1 P, 22 L (247-1) [USNM: BM; UCLA].
21. Aedes (Finlaya) heteropus Dyar
Figs. 5,42,43
1921. Aedes (Finlaya) heteropus Dyar 1921:152. TYPE: Lectotype 3 with genitalia slide
(1542), Alajuela, Costa Rica, 1 July 1921, bamboo, A. Alfaro [USNM, 24865; designa-
tion of Stone and Knight 1956:218].
Aedes (Finlaya) heteropus of Bonne and Bonne-Wepster (1925 :422); Dyar (1925b: 147).
Aedes (Finlaya) terrens in part of Lane (1939:105; 1953:686); Stone, Knight and Starcke (1959:
171); Belkin, Schick and Heinemann (1965:12); Forattini (1965 :395).
Aedes terrens in part of Kumm, Komp and Ruiz (1940:400,417).
Aedes (Finlaya) terrens var. podographicus of Edwards (1932:510); Knight and Marks (1952:549).
Aedes (Finlaya) podographicus in part of Dyar (1928:223).
Schick: Terrens Group of Aedes (Finlaya) 75
FEMALE. Head: Vertex with decumbent scales laterad of median longitudinal
line dark, narrow curved; occiput with erect scales pale; proboscis subequal in length
to or longer than femur I. Thorax (fig. 42): Complete acrostichal and posterior
dorsocentral lines absent; most posterior acrostichal seta at about 0.5 from anter-
ior end or in posterior half; fossal macula with mesal margin essentially straight,
supraalar macula not or narrowly joined to fossal macula; prescutellar space with
silver scales; midlobe of scutellum without silver scales; ssp scales present. Legs:
Femur I without knee spot; femora I and II wtih well-developed posterior patch
of silver scales in basal half, that of femur II larger; tarsus 1-I] with median dark
band complete, about 0.5; femur III with basal dark band incomplete or complete,
0.03, subapical dark band 0.36-0.37 (0.32-0.37); tarsus 1-III with basal silver band
0.13-0.14, apical silver band 0.19-0.25. Wing: Vein C with only a few basal silver
scales; vein R without silver scales.
MALE. Head: Vertex with all decumbent scales silver, narrow curved; palpus a-
bout 0.5-2 labellum lengths shorter than proboscis; segment 3 with several long apic-
al ventrolateral setae forming tuft as long as segments 4 and 5 combined; segment 4
with hairs of ventrolateral row usually long, somewhat sparse. Thorax (fig. 42): Mes-
onotal disc usually without strip of silver scales bordering fossal macula; complete
acrostichal and posterior dorsocentral lines present; fossal macula coextensive with
fossa. Legs: Tarsus 1-II with median dark band complete, about 0.33-0.5. Wing:
Vein C with basal silver scales usually forming small patch but sometimes developed
as interrupted line reaching crossvein h.
MALE GENITALIA (fig. 42). Sidepiece: Length 0.35 mm; basal tergomesal area
without dense patch of long setae; median sternomesal area with convexity and scler-
ite weakly developed, tuft absent. Prosophallus: Length 0.11 mm (0.13 mm in 1
specimen); mesal lobe with lateral portion inclined from between 15° and 30° to
about 30° from horizontal; stems essentially parallel or divergent; filament ratio
0.45-0.65; hook of filament not strongly angulate. Aedeagus: Length 0.12-0.13 mm.
PUPA (fig. 42). Cephalothorax: Hair 9-C single. Abdomen: Hair 1-I with primary
branches predominantly double-triple to triple-multiple; 2-II laterad of 3-II or mes-
ad for 0.3 or less the distance from 1-II to 3-II. Paddle: Apex not strongly pro-
duced; ventral midrib strongly pigmented for most length, weakly at apex.
LARVA (fig. 43). Head: Hair 5-C usually 5-branched (4-6); 6-C usually triple or 4-
branched (2-5); 7-C usually 5,6-branched; bmh usually branched but often single.
Thorax: Hair 1-P usually triple or 4-branched (3-5); 4-P usually triple (2-4); 5-P usu-
ally double or triple (2-4); 7-P usually double (1-3); 11-P about half length of 14-P;
8-T subequal in length to or longer than metathoracic pleural tubercle. Abdomen:
Hair 2-I usually triple or 4-branched (2-4); 3-III double. Segment VIIJ: Comb scales
32-53; free portion of midapical scale usually shorter than sessile portion; length
0.027-0.030 mm (0.022-0.035 mm). Siphon: Length 0.70-0.86 mm; L/S 2.13-2.22
(2.13-2.33); P/L 0.53-0.56 (0.50-0.57); H/L 0.61-0.65 (0.59-0.68). Anal Segment:
Ventral brush usually with 12 hairs (11-12); 4a-X usually 6-branched (4-6).
SYSTEMATICS. Aedes heteropus is distinguished from the other members of
the subgroup in the adults by a combination of the absence of complete mesono-
tal lines in the female and the presence of a complete acrostichal line in the male;
in the male genitalia by a combination of the absence of a median sternomesal
sclerite and tuft on the sidepiece and the moderately steeply inclined mesal lobe
of the prosophallus; and in the larva by a combination of the multiple hair 5-C
and the long 11-P.
The females of heteropus and homoeopus are very similar. Their separation in
76 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
the key, by the presence of silver scales on the midlobe of the scutellum in homoeo-
pus and the absence of the scales in heteropus, must be further verified. Probably
more reliable separation can be made on the basis of the pigmentation of the mid-
rib of the paddle if the pupa is available.
DISTRIBUTION (fig. 5). Costa Rica at elevations of 3100-3800 ft; 1 record,
based upon questionable determination, from El Salvador at an elevation between
1700 and 3300 ft. Material examined: 72 specimens; 22 6, 8 9, 1 pupa, 41 larvae; 1
individual rearing (larval).
COSTA RICA. Alajuela: Alajuela [elev. 3100 ft], 20 May 1921, bamboo, A. Alfaro, 2 d(1 4,
type series in part); May 1921, bamboo, 1 ¢, type series in part; June 1921, bamboo, A. Alfaro, 3
36, 1%; 1 July 1921, bamboo, A. Alfaro, 2 9, type series in part; 4 July 1921, bamboo, A. Alfaro,
3 6,3 2 (1 dG, type series in part); 30 July 1921, A. Alfaro, 1 6; 7 Oct 1921, A. Alfaro, 3 6; Oct
1921, A. Alfaro, 1 ¢ (1635), 4 6, 1 9; Apr 1922, A. Alfaro, 1 6, type series in part (USNM,
24865). San Jose: San Jose, Parque Bolivar, elev. 3800 ft, 8 June 1963, small treehole, height 7-8
ft, C. Hogue (CR 88), 1 Ip? (88-101), 2 6, 38 L (88-3) — small treehole adjacent to CR 88, height
7 ft, C. Hogue, 1 L (89-1) [UCLA]. San Jose, University of Costa Rica, elev. 3800 ft, 8 Nov 1962,
large treehole, J. Belkin, C-Hogue, W. Powder, 1 p? (CR 7-101).
EL SALVADOR. Cuscatlan: Cojutepeque [elev. 1700-3300 ft] , 1 6(626) [USNM].
Galindoi Subgroup
ADULTS. Head: Vertex of both sexes with decumbent scales along longitudinal
midline narrow; occiput of both sexes with erect scales pale; proboscis of female
longer than femur I; palpal segment 3 of males with | or 2 short to moderately long
hairs; segment 4 with hairs of ventrolateral row short to moderately long, sparse.
Thorax: Mesonotal disc of both sexes not transversely silvered; complete acrostichal
or posterior dorsocentral lines absent in females; acrostichal setae present; fossal
macula of females not coextensive with fossa, variously reduced; of males coextensive
with fossa or reduced mesally; supraalar macula not truncate posteriorly, in females
not joined to or narrowly to broadly joined to fossal macula; ppn silver scaled; ssp
scales absent; pra hairs usually pale; upper stp and mep scale patches not contiguous.
Legs: Mid- and hindlegs not shaggy; mid- and hindtarsi with conspicuous silver bands;
tarsus I-I with or without narrow apical silver band; femur II with knee spot absent
or narrow, a single row at apex of segment; tarsus 1-II with median dark band com-
plete, usually very broad; tarsus 2-II with complete apical dark band; tarsus 5-II
without silver scales; femur III with basal dark band incomplete or complete, narrow
or broad; tarsus 5-III without silver scales. Wing: Vein C of both sexes with small
basal patch of silver scales; veins R and Cu of males without silver scales.
MALE GENITALIA. Sidepiece: Basal tergomesal area with dense patch of long
setae; median sternomesal area with or without strongly developed sclerite, with tuft;
specialized subapical sternal seta absent. Prosophallus: Mesal lobe with lateral por-
tion inclined from between 15° and 30° to 45° from the horizontal; stem usually
not bowed; filament with hook strongly angulate.
PUPAE. Cephalothorax: Without pale inverted V-shaped marking; hair 5-C less
than twice length of 4-C; 9-C usually single. Abdomen: Hair 1-I with primary branch-
es predominantly double to multiple; 2-II usually laterad to just barely mesad of 3-
II; 3-III single. Paddle: Pigmented; apex not or weakly produced; hair 1-P shorter
than paddle.
LARVAE. Head: Hair 5-C usually multiple, sometimes triple; 6-C usually double
to multiple; 7-C more than half length of 6-C; 11-C longer than mental plate; 14-C
Schick: Terrens Group of Aedes (Finlaya) qT
longer than mental plate, usually branched; bmh usually branched. Antenna: Hair 1-
A single to multiple. Thorax: Hair 11-P at least half length of 14-P; 14-P branched; 3-
M single; 4-M single to multiple; 8-T shorter, but more than half length of, or longer
than metathoracic pleural tubercle. Abdomen: Hair 5-I at least subequal in length to
4-I; 2-II slightly mesad of (about 1 alveolus width) to laterad of 4-II; 3-III single or
double; 10-III single; 3-VI usually single; 4-VII single; 5-VII caudad of 4-VII, more
than half length of 3-VII; 10-VII single to triple; 12-VII single. Segment VII: Mid-
apical comb scale with free portion subequal in length to or longer than sessile por-
tion; 2-VIII single. Anal Segment: Saddle extending around segment for moderate
distance, ventral margin without deep slit.
DISCUSSION. The Galindoi Subgroup is characterized in the adults by a com-
bination of the absence of complete silver lines on the mesonotal disc of both
sexes, the presence of acrostichal setae, the absence of a scale patch on the ssp,
the absence usually of a knee spot on femur II and the very broad median dark band
of tarsus 1-II; and in the larvae by the position of hair 2-II, which is situated farther
laterad than in the other groups. At least 2 of the 3 species show hairy-nonhairy
dimorphism in the larvae. The male genitalia and pupa do not show characters that
would differentiate them from the other subgroups.
Three species comprise the subgroup, galindoi, campana and daryi. Aedes galindoi
and campana are closely related; the former occurs in the Canal Zone and the latter
in the highlands of central Panama. Aedes daryi is represented by 2 widely disjunct
montane populations, 1 in eastern Panama and 1 near the Atlantic coast of Guate-
mala.
22. Aedes (Finlaya) galindoi Schick, n.sp.
Figs. 5,44,45
TYPE: Holotype 6 (207C-20) with associated pupal and larval skins (5-147) and genitalia slide
(672113-4), Barro Colorado Island [elev. 100-500 ft], Canal Zone, Panama, 15 May 1945, tree-
hole, W.H.W. Komp [USNM]. Allotype 2 (207C-26) with 2 sets of pupal and larval skins, 1 set
representing actual associated stages (5-152), same data as holotype [USNM]. Paratypes: 2 lpd
(207C-25, 5-139; 207C-41, 5-148), 1 Ip? (207C-26, 5-152; see allotype), 1 1 (207C-1), 1 L, same
data as holotype [BM; UCLA]. This species is dedicated to Pedro Galindo of the Gorgas Memorial
Laboratory, Panama.
FEMALE. Head: Vertex with decumbent scales laterad of median longitudinal
line dark, broad. Thorax (fig. 44): Most posterior acrostichal seta usually at about
0.5 from anterior end (between 0.33 and 0.5 in | specimen); fossal macula reduced
anteriorly and mesally; supraalar macula reduced in size, disjunct from fossal mac-
ula; prescutellar space and midlobe of scutellum without silver scales; pra hairs pale
or dark. Legs: Femora I and II without posterior patch of silver scales; femur II
without knee spot; tarsus 1-I] with median dark band from slightly greater than 0.5
to about 0.67; femur III with basal dark band 0.12-0.16 (0.07-0.16), subapical dark
band 0.28-0.34; tarsus 1-III with basal silver band 0.10-0.14, apical silver band 0.09-
0.16. Wing: Vein R without silver scales.
MALE. Head: Vertex wtih decumbent scales laterad of median longitudinal line
dark or silver, broad; palpus about 1-2 labellum lengths shorter than proboscis; seg-
ment 3 with | or 2 short hairs at ventrolateral apex (hairs apparently absent in |
78 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
specimen); segment 4 with hairs of ventrolateral row short, sparse. Thorax (fig. 44):
Fossal macula not reaching mesal margin of fossa; pra hairs pale. Legs: Femur II
without knee spot; tarsus 1-IJ with median dark band as in female.
MALE GENITALIA (fig. 44). Sidepiece: Length 0.36-0.39 mm; median sterno-
mesal area with convexity weakly to moderately developed, sclerite well developed,
tuft very dense, the setae wavy. Prosophallus: Length 0.11-0.12 mm; median lobe
projecting cephalad of lateral lobe; mesal lobe with lateral portion inclined at about
45° from horizontal; stems usually divergent or essentially parallel (sometimes one
stem bowed, converging towards the other); filament ratio 0.55-0.80. Aedeagus:
Length 0.13 mm (0.12-0.13 mm).
PUPA (fig. 44). Cephalothorax: Hair 9-C usually single, infrequently double. Ab-
domen: Hair 1-I with primary branches usually predominantly double to triple,
sometimes triple-multiple; 2-II laterad of 3-II] or mesad for less than 0.3 the dis-
tance from 1-II to 3-II. Paddle: Ventral midrib usually weakly pigmented, sometimes
moderately pigmented for most length, weakly at apex.
LARVA (fig. 45). Head: Hair 5-C usually 5-7 branched (3-7); 6-C most usually 4-
branched (1-6); 7-C usually 5-7 branched (4-9); 14-C usually double, infrequently
single or triple; bmh usually double, infrequently single. Antenna: Hair 1-A usually
single, infrequently double. Thorax: Hair 1-P usually 4-branched (3-5); 4-P usually
triple or 4-branched (2-4); 5-P usually double or triple (1-5); 7-P usually double to 4-
branched (2-6); 4-M usually double, infrequently triple; 8-T longer than metathora-
cic pleural tubercle. Abdomen: Hair 3-III double; 3-VI single; 10-VII usually single
(1-2). Segment VIII: Comb scales 35-47; free portion of midapical scale longer than
sessile portion, length 0.034-0.040 mm. Siphon: Length 0.71-0.85 mm; L/S 2.18-
2.27; P/L 0.57-0.59; H/L 0.63-0.67. Anal Segment: Ventral brush usually with 12
hairs (11-13); 4a-X usually 5 ,6-branched (5-7).
SYSTEMATICS. Aedes galindoi and campana are very closely related. The fol-
lowing similarities set them apart from the other species of the subgroup, daryi,
fossal and supraalar maculae of females greatly reduced, posterior surface of femora
1-I] without silver scales, median sternomesal tuft of the sidepiece very dense and
unique in consisting of wavy hairs, and hair 1-A of the larvae usually single.
Aedes galindoi differs from campana in the usually shorter sidepiece, 0.36-0.39
mm vs. 0.39-0.41 mm, the more strongly produced median lobe of the prosophallus,
the greater filament ratio, 0.55-0.80 vs. 0.45-0.55, the greater number of comb
scales, 35-47 vs. 23-32, the scales arranged in 4 rows rather than 3, the relatively
longer free portion of the apical scale and the greater L/S, 2.18-2.27 vs. 1.96-2.08.
Aedes galindoi is found at low elevations (0-100 ft) and campana at high elevations
(2800 ft).
The 2 larvae of PA 856 (Barro Colorado Island) differ from the others in the
fewer number of branches of some of the hairs. Hair 5-C is single or double in
these specimens but at least 5-branched in the other lots and 6,7-C, 7,8-P,T and
1-X show at least 2 fewer branches than in the other lots.
DISTRIBUTION (fig. 5). Canal Zone, Panama. Material examined: 52 specimens;
11 6, 8 2, 16 pupae, 17 larvae; 13 individual rearings (8 larval, 4 pupal, 1 incom-
plete). For the Komp collections see the explanatory chapter.
PANAMA. Canal Zone: Barro Colorado Island [elev. 100-500 ft], 15 May 1945, treehole, W.
Komp (type series), 3 lpd (207C-20,25,41), 2 9, 2 p, 2 1 (207C-26), 1 1(207C-1), 1 L [USNM;
BM; UCLA] — 1 Ipd (207C-19); 22 May 1945, deep treehole at ground level, W. Komp, 2 Ip?
(207A-16,17), 1 L, 1 1 (5-256, adult number unknown); 26 June 1945, W. Komp (adult numbers
unknown), | Ip (5-146), 11(5-451) [UCLA; USNM] ; 26 July 1935, L. Rozeboom, 1 6 (PAR 81-2)
Schick: Terrens Group of Aedes (Finlaya) 79
[UCLA]; 3 Dec 1965, treehole, height 3 ft, A. Quinonez (PA 856), 2 lp? (856-12,13), 2 6, 1 p
(856-4) [UCLA]. Fort Sherman [elev. 0-100 ft], 9 Sept 1949, 1 d [UCLA]. Margarita [elev. 0-
100 ft], 21 Jan 1922, J. Shropshire, 1 9 [USNM]. Mojinga Swamp [elev. near sea level] , 13 Dec
1964, treehole (PA 721), 3 pd (721-13,102,103), 1 p? (721-12), 1 L(721-2) [UCLA].
23. Aedes (Finlaya) campana Schick, n.sp.
Figs. 5 ,46,47
TYPE: Holotype 3 (PA 518-109) with associated pupal and larval skins, Cerro Campana [elev.
2800 ft], Panama, Panama, 16 Aug 1943, wood bowl [USNM]. Allotype 2 in alcohol (PA 518-
172) with associated pupal and larval skins, same data as holotype [USNM]. Paratypes: 1 lpdé
(518-121), 1 L (518-2), same data as holotype [UCLA].
?Aedes (Finlaya) terrens in part of Galindo, Carpenter and Trapido (1951:121; 1955:169); Trap-
ido, Galindo and Carpenter (1955a:530).
FEMALE. Essentially as in galindoi, but most posterior acrostichal seta in poster-
ior half in all specimens; pra hairs pale. Legs: Femur III with basal dark band 0.11,
subapical band 0.28; tarsus 1-III with basal silver band 0.13, apical silver band 0.14.
MALE. Essentially as in galindoi, but femur II with narrow knee spot in 1 of 2
specimens.
MALE GENITALIA (fig. 46). Sidepiece: Length 0.39-0.41 mm; median sterno-
mesal area with convexity moderately developed, sclerite well developed, tuft very
dense, the setae wavy. Prosophallus: Length 0.11-0.12 mm; median lobe not pro-
jecting cephalad of lateral lobe; mesal lobe with lateral portion inclined at about
45° from horizontal; stems divergent; filament ratio 0.55-0.80. Aedeagus: Length
0.13 mm. |
PUPA (fig. 46). Cephalothorax: Hair 9-C sitgle. Abdomen: Hair 1-I with primary
branches predominantly multiple; 2-II laterad of or just barely mesad of 3-II. Pad-
dle: Ventral midrib weakly pigmented.
LARVA (fig. 47). Head: Hair 5-C usually 5-branched (4-6); 6-C double or triple;
7-C 4-6 branched; 14-C double or triple; bmh double. Antenna: Hair 1-A single.
Thorax: Hair 1-P triple or 4-branched; 4,5-P double or 4-branched; 7-P triple; 4-M
double; 8-T longer than metathoracic pleural tubercle. Abdomen: Hair 3-III single or
double; 3-VI single; 10-VII single. Segment VIII: Comb scales 23 (28 on other side
of this specimen)-32; free portion of midapical scale subequal in length to or slightly
shorter than sessile portion, length 0.035 mm. Siphon: Length 0.80-0.85 mm; L/S
1.96-2.08; P/L 0.55-0.58; H/L 0.64-0.66. Anal Segment: Ventral brush with 11 or
12 hairs; 4a-X 5-branched.
SYSTEMATICS. See galindoi.
DISTRIBUTION (fig. 5). Provinces of Cocle and Panama, Panama, at elevations
between 1800 and 3300 ft. Material examined: 10 specimens; 2 6, 2 9, 3 pupae; 3
larvae; 3 individual rearings (larval).
PANAMA. Cocle: El Valle [elev. 1800-3300 ft], Nov 1946, N. Krauss, 1 9 [USNM]. Panama:
Cerro Campana [elev. 2800 ft], 16 Aug 1953, wood bowl (PA 518, type series), 2 Ipd (518-109,
121), 1 lp? (518-172), 1 L (518-2) [USNM; UCLA].
80 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
24. Aedes (Finlaya) daryi Schick, n.sp.
Figs. 548-51
TYPE: Holotype 2 (GUA 33-31) with associated pupal and larval skins, Finca Trece Aguas,
nearest town Senahu, elev. 2800 ft, Alta Vera Paz, Guatemala, 7 July 1964, cut or broken bam-
boo, T. and J. Zavortink [USNM]. Allotype 36 (GUA 33-110) with associated pupal skin, same
data as holotype [USNM]. Paratypes: 1 pd (33-44), 5 L (33-4), same data as holotype [UCLA].
This species is dedicated to Mario Dary R. of the Universidad de San Carlos, Guatemala.
FEMALE. Head: Vertex with decumbent scales laterad of median longitudinal
line dark, broad. Thorax (figs. 48,50): Most posterior acrostichal seta usually at
about 0.5 from anterior end or in posterior half of line; fossal macula not reduced
anteriorly, broader in Trece Aguas, Guatemala, than in Cerro Mali, Panama (see figs.
48,50); supraalar macula usually broadly joined to fossal macula (narrowly joined
in PA 381-102); prescutellar space and midlobe of scutellum with silver scales; pra
hairs pale. Legs: Femora I and II with well-developed posterior patch of silver scales
in basal half, that of femur II larger; tarsus 1-IJ with median dark band about 0.7;
femur III with basal dark band incomplete, subapical dark band 0.23-0.33 (in Trece
Aguas, 0.23; in Cerro Mali, 0.25-0.33); tarsus 1-III with basal silver band 0.08-0.13
(in Trece Aguas, 0.08: in Cerro Mali, 0.10-0.13); apical silver band 0.07-0.12. Wing:
Vein R without silver scales.
MALE. Head: Vertex wtih decumbent scales laterad of median longitudinal line
silver, broad; palpus about 3 labellum lengths shorter than proboscis; segment 3 with
1 or 2 moderately long apical ventrolateral hairs, not as long as segments 4 and 5
combined: segment 4 with hairs of ventrolateral row short to moderately long,
sparse. Thorax (figs. 48,50): Fossal macula coextensive with fossa; pra hairs pale.
Legs: Femur II without knee spot; tarsus 1-I] with median dark band as in female.
MALE GENITALIA (figs. 48,50). Sidepiece: Length 0.35-0.42 mm; median stern-
omesal area with convexity absent or weakly developed, sclerite weakly developed,
e@fuft present but not strongly differentiated, the setae not wavy. Prosophallus: Length
O.11-0.13 mm; median lobe not projecting cephalad of lateral lobe; mesal lobe with
lateral portion inclined between 15° and 30° from horizontal; stems convergent;
filament ratio 0.50-0.70. Aedeagus: Length 0.13-0.14 mm.
PUPA (figs. 48,50). Cephalothorax: Hair 9-C single. Abdomen: Hair 1-I with pri-
mary branches predominantly double to multiple: 2-II always laterad of 3-II. Pad-
dle: Ventral midrib weakly to moderately pigmented, not more strongly at apex.
LARVA (figs. 49,51). Head: Hair 5-C usually 6-8 branched (5-10); 6-C double to
10-branched; 7-C 6-11 branched; 14-C usually triple (2-5); bmh double to 4-branch-
ed. Antenna: Hair 1-A single to 4-branched. Thorax: Hair 1-P most usually 4-6
branched (2-6); 4-P double to 8-branched; 5-P usually 3-5 branched (2-6); 7-P usually
triple (2-5): 8-T subequal in length to or longer than metathoracic pleural tubercle.
Abdomen: Hair 3-III usually double (1-3); 3-VI usually single, infrequently double;
10-VITI usually single or double (1-3). Segment VIII: Comb scales 18-38; free portion
of midapical scale subequal in length to or longer than sessile portion, length 0.035-
0.046 mm (0.033-0.047 mm). Siphon: Length 0.82-0.94 mm; L/S 2.27-2.70; P/L
O.51-0.58 [in Cerro Mali, Panama 0.51-0.57; in Trece Aguas, Guatemala 0.56-0.58 ] ;
H/L 0.57-0.67 [in Cerro Mali 0.57-0.63; in Trece Aguas 0.65-0.67]. Anal Segment:
Ventral brush usually with 12 hairs (11-13): 4a-X usually 5-branched (4-6).
SYSTEMATICS. Aedes daryi differs from galindoi and campana in the more ex-
Schick: Terrens Group of Aedes (Finlaya) 81
tensively developed fossal and supraalar maculae of the female, the presence of a
patch of silver scales on the posterior surface of femora I and II, the more poorly
developed sternomesal tuft of the sidepiece of the male genitalia, and the usually
branched hair 1-A of the larva. The larval comb scales are exceptionally large for
the Terrens Group.
Aedes daryi is known from 2 widely separated areas, Trece Aguas, Guatemala
(GUA 33), and Cerro Mali in eastern Panama (PA 349,367,381). The only appar-
ently significant difference I can find between the populations from these areas is
the broader fossal macula of the females from Trece Aguas.
The larvae show dimorphism in hairiness. The larvae of the Guatemalan and PA
381 collections are hairy and those of PA 349 and PA 367 are nonhairy. The most
striking difference between these 2 forms is in the number of branches of hair 1 of
the abdominal segments. Of greater significance, however, is the larger number of
branches in the hairy forms of some of the hairs whose branching provides import-
ant subgroup characters in the Terrens Group, hair 1-A is single to double in the
nonhairy form vs. double to 4-branched in the hairy form; 4-M single or double
vs. triple or 4-branched; and 10-VII single vs. single to triple. Also, hair 8-T is sub-
equal in length to the metathoracic pleural tubercle in the nonhairy form vs. longer
than the tubercle in the hairy form (see also gabriel).
DISTRIBUTION (fig. 5). Department of Alta Vera Paz, Guatemala, at elevation
of 2800 ft; province of Darien, Panama, at elevations of 4500-4900 ft. Material ex-
amined: 30 specimens; 3 6, 5 9, 8 pupae, 14 larvae; 8 individual rearings (6 larval,
2 pupal).
GUATEMALA. Alta Vera Paz: Finca Trece Aguas, elev. 2800 ft, 7 July 1964, cut or brok-
en bamboo, T. and J. Zavortink (GUA 33, type series), 1 lp? (33-31), 2 pd (33-110,44), 5 L (33-4)
[USNM; UCLA].
PANAMA. Darien: Cerro Mali [8° 07’ N - 77° 14’ W], auxiliary ridge W of, elev. 4900 ft, 22
May 1963, treehole, height 20 ft (PA 349), 2 lp? (349-101,102), 1 L (349-2); western slopes, elev.
4700 ft, 28 May 1963, treehole, height 10 ft, 1 lp¢ (MEX 367-101); 4 mi W summit, elev. 4500 ft,
8 June 1963, treehole, height 12 ft (PA 381), 2 lp? (381-101,102), 2 L (381-1) [UCLA].
Podographicus Subgroup
ADULTS. Head: Vertex of both sexes with decumbent scales along longitudinal
midline narrow; occiput of both sexes with erect scales pale; proboscis of female
shorter than or subequal in length to femur I; palpal segment 3 of males with 1-3
apical ventrolateral hairs of moderate length, shorter than segments 4 and 5 com-
bined; segment 4 with hairs of ventrolateral row of moderate length, sparse. Thor-
ax: Mesonotal disc of both sexes not transversely silvered; complete acrostichal or
posterior dorsocentral lines absent in female; acrostichal setae absent; fossal macu-
la of both sexes not coextensive with fossa, reduced mesally; supraalar macula not
truncate posteriorly, in females disjunct from fossal macula; ppn silver scaled; ssp
scales present or absent; pra hairs of female pale; upper stp and mep scale patches
not contiguous. Legs: Mid- and hindlegs not shaggy; mid- and hindtarsi with con-
spicuous silver bands; tarsus 1-I with or without narrow apical silver band; femur II
with knee spot moderately broad, the scales extending to about level of anterior sub-
apical setae; tarsus 1-IJ with median dark band incomplete or complete, narrow; tar-
sus 2-II with incomplete or complete apical dark band; tarsus 5-II without silver
scales; femur III with basal dark band incomplete or complete, narrow; tarsus 5-II]
82 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
without silver scales. Wing: Vein C of females with small basal patch or line of silver
scales reaching about 0.5 to crossvein h, of males usually with a line reaching about
0.5 to h; vein R of males without silver scales or with basal line of silver scales reach-
ing level of crossvein h; vein Cu of males without silver scales.
MALE GENITALIA. Sidepiece: Basal tergomesal area without dense patch of
long setae; median sternomesal area without strongly developed sclerite and with-
out tuft; specialized subapical sternal seta absent. Prosophallus: Mesal lobe with lat-
eral portion usually inclined between 15° and 30° from horizontal; stems usually
bilaterally bowed and convergent; filament with hook not strongly angulate.
PUPA. Cephalothorax: Without pale inverted V-shaped marking; hair 5-C less than
twice length of 4-C; 9-C single. Abdomen: Hair 1-I with primary branches predom-
inantly single to multiple; 2-I] mesad of 3-II usually for 0.3 or more the distance
from 1-II to 3-II; 3-III single. Paddle: Pigmented; apex usually produced, some-
times strongly so; hair 1-P shorter than paddle.
LARVA. Head: Hairs 5,6-C usually single; 7-C more than half length of 6-C; 11-
C longer than mental plate; 14-C longer than mental plate, usually single; bmh usual-
ly single. Antenna: Hair 1-A single. Thorax: Hair 11-P less than half length of 14-P;
14-P usually single; 3-M single; 4-M double; 8-T more than half length of or subequal
in length to metathoracic pleural tubercle. Abdomen: Hair 5-I at least subequal in
length to 4-I; 2-II well mesad of 4-II; 3-III double; 10-III single; 3-VI single; 4-VII us-
ually single; 5-VII caudad of 4-VII, usually less than half or about half length of 3-
VII; 10,12-VII single. Segment VIII: Midapical comb scale with free portion shorter
or longer than sessile portion; 2-VIII single. Anal Segment: Saddle extending around
segment for moderate distance, ventral margin without slit.
DISCUSSION. The Podographicus Subgroup is characterized in the adults by a
combination of the short proboscis of the female, the absence of silver markings
on the mesonotal disc of both sexes, the absence of acrostichal setae, the disjunct
fossal and supraalar maculae of the female and the incomplete or narrow basal
dark band of femur III; and in the male genitalia by the inwardly bowed. stems of
the prosophallus. The larva shows nonmultiple hairs 5,6-C and cannot be readily
separated from the larvae of the species of the Metoecopus, Homoeopus, and Heter-
opus Subgroups that show this combination of characters. The pupae do not show
distinctive subgroup characters.
The subgroup is known from coastal Mexico and Central America and from the
Maracay area of Venezuela.
Only 1 species, podographicus, is formally recognized here but there is a popula-
tion from outside the range of typical podographicus which may represent a sibling
species (see podographicus).
25. Aedes (Finlaya) podographicus Dyar & Knab
Figs. 6,52-57
1906. Aedes podographicus Dyar and Knab, 1906:165. TYPE: Lectotype ? (325j), Sonsonate,
FE] Salvador [USNM, 10015; designation of Stone and Knight 1956:224].
Aedes (Finlaya) podographicus of Dyar (1921:151).
Aedes (Finlaya) podographicus in part of Dyar (1928:223-224); Martini (1935:56).
Aedes podographicus of Theobald (1910:484); Dyar (1918:73,80).
Aedes (Finlaya) terrens podographicus of Horsfall (1955 :699).
Schick: Terrens Group of Aedes (Finlaya) 83
Aedes (Finlaya) terrens var. podographicus in part of Edwards (1932:150); Knight and Marks
&. (1 952:549).
Aedes (Finlaya) terrens in part of Lane (1939:105; 1953:686-687); Trapido, Galindo and Car-
penter (1955a:550); Stone, Knight and Starcke (1959:171); Belkin, Schick and Heinemann
(1965:22); Forattini (1965 :395).
? Aedes (Gualteria) terrens in part of Diaz Najera (1963:131; 1966:61).
Aedes terrens in part of Kumm, Komp and Ruiz (1940:400 416); Kumm and Zuniga (1942:407).
Aedes insolitus in part of Dyar and Knab (1906b:203).
FEMALE. Head: Vertex usually with decumbent scales of area just adjacent to
median longitudinal line dark, narrow curved, and scales laterad of these dark,
broad (in state of Campeche, Mexico, scales laterad of line generally dark, broad,
some silver scales scattered among these). Thorax (fig. 52): Supraalar macula not or
rarely narrowly joined to fossal macula; ssp scales variable in occurrence [in Panama
present; in Costa Rica, and state of Campeche, Mexico, present or absent; in Nicar-
agua, El Salvador, Guatemala and San Blas, Mexico, usually absent]. Legs: Femora
I and II with well-developed posterior patch of silver scales in basal half, subequal in
size; median dark band of tarsus 1-II usually incomplete or complete and width less
than 0.33, infrequently complete, as broad as about 0.33; tarsus 2-II usually with |
complete dark apical band but sometimes silvered along entire length on one side;
femur III with basal dark band incomplete or complete, as broad as 0.05 (0.10),
subapical dark band 0.33-0.45 [in San Blas 0.33-0.40; on Pacific side of Central
America 0.37-0.42 (0.33-0.45); in state of Campeche 0.45]; tarsus 1-III with basal
silver band when present as broad as 0.10 [in most localities when present as broad
as 0.08 but usually less; in state of Campeche present, 0.10]; apical silver band
0.20-0.38 [in most localities 0.24-0.33 (0.20-0.35); in state of Campeche, 0.38].
Wing: Vein C with basal silver scales forming small patch or line reaching about
0.5 to crossvein 4; vein R usually without or with only a few basal silver scales (in
San Blas with basal patch of scales).
MALE. Head: Vertex with decumbent scales laterad of median longitudinal line
dark or silver, otherwise as in female; palpus subequal in length to proboscis to
about | labellum length shorter. Thorax (fig. 52). Legs: Tarsus 1-I] with median
dark band as in female except in state of Campeche, Mexico, where band complete,
0.6. Wing: Vein C with basal silver scales usually forming line reaching about 0.5
to crossvein h, occasionally with scales forming small patch; vein R with silver scala-
tion variably developed [on Pacific side of Central America and in state of Cam-
peche, Mexico, absent; in San Blas, Mexico, present, forming line reaching level of
crossvein h].
MALE GENITALIA (figs. 52,54,56). Sidepiece: Length 0.35-0.39 mm (0.34-0.40
mm). Prosophallus: Length 0.12-0.14 mm (0.11-0.15 mm); mesal lobe with lateral
portion usually moderately inclined, between 15° and 30° from the horizontal (in-
frequently somewhat less than 15° or as much as about 30°); filament ratio 0.40-
0.50 (0.30-0.65). Aedeagus: Length 0.13-0.14 mm (0.13-0.15 mm).
PUPA (figs. 52,54,56). Abdomen: Hair 1-I with primary branches usually predom-
inantly double-triple to multiple, infrequently predominantly single; 2-II in line with
3-II or mesad of 3-II for more than 0.3 distance from 1-II to 3-II [in Panama, San
Blas, Mexico, and state of Campeche, Mexico, usually 0.3 or more mesad; in other
localities 0.4 or more]. Paddle: Apex usually weakly produced in most localit-
ies, moderately to strongly produced in San Blas; ventral midrib weakly or mod-
erately pigmented for most length, weakly at apex.
LARVA (figs. 53,55,57). Head: Hair 5-C usually single, rarely double; 6-C single;
84 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
7-C usually triple (2-4); 14-C, bmh usually single, very rarely double. Thorax: Hair
1-P usually double or triple (1-5); 4-P usually double (1-4); 5-P single or double; 7-P
usually double (1-4). Abdomen: Hair 3-VI single, rarely double; 4-VII usually single,
rarely double. Segment VIII: Comb scales 46 to more than 100; length of free por-
tion of midapical comb scale 0.019-0.035 mm [in San Blas, Mexico, 0.019-0.025
mm; in state of Campeche, Mexico, 0.025-0.026 mm; on Pacific side of Central
America, 0.026-0.032 mm (0.024-0.035 mm)]. Siphon: Length very variable, 0.71-
1.05 mm, most lower values from Nicaragua; L/S 2.13-2.56; P/L 0.55-0.59 (0.50-
0.62); H/L 0.60-0.69. Anal Segment: Ventral brush usually with 12 hairs, less usu-
ally 13, in San Blas 11 or 12 (11-14); 4a-X usually with 7 or 8 branches, in state of
Campeche, 6,7-branched (6-11).
SYSTEMATICS. Typical podographicus occurs in the Pacific lowlands of Central
America from Tehuantepec, Mexico, south into Panama. There are collections at
hand from 3 coastal lowland areas beyond this range, San Blas in the state of Nay-
arit, Mexico; near Champoton in the state of Campeche, Mexico; and in British Hon-
duras. The Mexican specimens deviate from typical podographicus as follows. Those
from San Blas show a more extensive silver scalation at the base of wing R, a pupal
paddle which is more strongly produced apically and a shorter free portion of the
apical comb scale of the larva; those from the state of Campeche show a broader
median dark band of tarsus 1-II in the male and broader silver bands of tarsus 1-[I in
the female (the length of the free portion of the apical comb scale falls into the up-
per end of the range of the San Blas specimens and the lower end of the range of
typical podographicus). The specimens from British Honduras are females taken in
light traps and are in too poor a condition to be treated in any detail; the legs bands
of tarsus 1-III, however, are not broad as in the females from the nearby state of
Campeche, the widths falling into the range of typical podographicus.
In the Maracay area of Venezuela, which is of moderately high elevation, there
is another podographicus-like form known only by the adults. This population is
indistinguishable from the typical podographicus of Central America and shows the
same morphological variation except that the silver scalation at the base of vein R is
more variable and may be developed to the same extreme as in the San Blas popu-
lation.
I am not describing at least the San Blas and Campeche populations as distinct
species because of (1) the few differences between these and the typical populations
of podographicus, (2) the uncertainty regarding the taxonomic significance of 3 of
the 4 characters involved, the silver scalation at the base of vein R which is so vari-
able in the Maracay area, the produced apex of the paddle which may be very vari-
able in other species (e.g. vargasi), and the broad tarsal bands which are not unusu-
ally broad in the specimens of the opposite sex from the area in question, and (3)
the lack of sufficient material from the coastal areas beyond the range of typical
podographicus to ascertain the variability of the characters and whether clinal vari-
ations are involved. I think that these 2 populations will be shown to represent at
best ‘“‘forms’’ of podographicus (or subspecies, depending upon the taxonomist’s
philosophy) when the fauna of the area becomes more adequately known. Conse-
quently, I am treating these as populations of podographicus here. The population
from Maracay area can only be treated as ‘“podographicus complex” until the im-
mature stages can be studied.
Aedes podographicus of the Pacific coast shows marked variation in the develop-
ment of the ssp scales, the branching of some larval hairs and in some of the siphon
indices. The development of a ssp scale patch shows a clinal variation, being present
Schick: Terrens Group of Aedes (Finlaya) 85
in all specimens in the southern part of the range and usually absent in those in the
northern part of the range (see description). Clinal variations in the branching of
hairs 1,4,5-P (see description) and 1-II-VII and 6-III-VI are not apparent. Hairs 1-
II-VII are often single in El Salvador and Panama; usually double or triple in Nicar-
agua and San Blas; and usually triple or multiple in Guatemala. Hairs 6-III-VI are
usually single in El Salvador; single or double in Panama; and usually double in
other localities. The L/S and H/S siphon indices also show geographical variations
of a nonclinal nature (see description).
DISTRIBUTION (fig. 6). Coastal lowlands of Mexico and Central America. Mater-
ial examined: 635 specimens; 96 6, 130 2, 157 pupae, 252 larvae; 106 individual
rearings (49 larval, 52 pupal, 5 incomplete.
COSTA RICA. Alajuela: San Mateo [elev. 800 ft], 1 6 (583) [USNM] . Guanacaste: Samara,
Nicoya Peninsula, elev. near sea level, 23 Aug 1964, small treehole, C. Hogue, Miranda (CR 196,
197), 8 L (196-2), 1 L (197-2) [UCLA] . Puntarenas: El Roble (near Puntarenas), [elev. 0-300 ft] ,
16 June 1963, T. Aitken, 1 d (207B-12) [UCLA]. Esparta [elev. 700 ft], 7 June 1943, T. Aitken,
5 6, 4 2; 1943, T. Aitken, 1 6, 1 9 [USNM]. Macacona [elev. 300-1700 ft] , 28 May 1943, tree-
hole, T. Aitken, 4 2 (207B-6); 1 June 1943, treeholes, T. Aitken, 2 6, 6 9? (2073-4) [UCLA]. Rio
Aranjuez, ? 9 Sept 1905, F. Knab (340), 2 6 (340a), 1 6, 1 9 (340b), 1 2 (340c) [USNM].
EL SALVADOR. San Miguel: San Miguel [elev. 300-700 ft] , ? May 1931, 9 6, 3 9 (207B-43), 1
? (531-D) [USNM]. Sonsonate: Canton El Castano, nearest town Sonsonate (elev. 730 ft), 1
Aug 1964, large treehole, height 12 ft, A. Quinonez (SAL 1), 4 Ip? (1-11,13,18,19), 2 pd (1-101,
ft (SAL 2), 1 Ip? (2-12), 1 P, 12 L (2-1) [UCLA]. Finca San Dionisio, nearest town San Antonio,
4 Aug 1964, large treehole, height 6 ft, A. Quinonez (SAL 8), 1 p? (8-100), 1 P, 25 L (8-2) — cut or
broken bamboo (SAL 10), 1 Ipd (10-21), 4 P, 2 L (10-2) [UCLA]. Sonsonate [elev. 700-1600 ft] ,
18 Aug 1905, treehole, F. Knab (325, lectotype collection), 4 6 (325c,e,f,h), 5 6 (325j) [USNM].
Usultan: San Juan del Gozo (Peninsula de) [elev. 0-300 ft] , H. Kumm, 2 6 (207A-28) [UCLA].
GUATEMALA. Escuintla: Escuintla, vicinity (63.8 km S Guatemala), elev. ca 660 ft, 10
July 1964, large treehole, height 10 ft, J. and T. Zavortink (GUA 39), 1 Ipd (39-33), 7 lp? (39-30-
32,34-36,38), 1 pd (39-104), 1 IP (39-33), 14 6, 14 9, 3 P, 35 p, 20 L, 3 1 (39-3). Escuintla,
vicinity (52.5 km S Guatemala), elev. 660-730 ft, 10 July 1964, large treehole, height 6 ft, T.
and J. Zavortink (GUA 40), 1 Ipd (40-11), 3 lp? (40-10,12,13), 5 pd (40-100-102,104,105), 1 p?
(40-103), 1 L (40-1); 10 July 1964, large treehole, height 6 ft, P. Cowsill (GUA 41), 3 lp? (41-10-
13), 5 pd (41-100-104), 1 p? (41-105), 3 P, 25 L (41-1) [UCLA]. San Jose, N of (87.5 km from
Guatemala on hwy CA 9), elev. near sea level, 10 July 1964, small treehole, height 6 ft, T. and J.
Zavortink (GUA 38), 1 Ipd (38-11), 2 Ip? (38-10,12), 1 p? (38-100), 5 L (38-1) [UCLA]. San
Jose de Guatemala [probably Department of Escuintla and elev. near sea level], 15 July 1943,
treehole, D. Hall, 1 6, 3 2 (207A-37); 15 July 1943, D. Hall, 2 9, 2 p (207A-43); 20 July 1943,
D. Hall, 2 2, 2 p, 1 1(207D-10); July 1943, D. Hall, 1 9, 1 1(207D-11) [USNM; UCLA]. Retalhu-
leu: Near San Sebastian, 50 yds E Rio Mulua bridge, elev. 990 ft, 2 July 1964, cut or broken bam-
boo, P. Cowsill, T. Zavortink (GUA 29), 1 Ipd (29-11), 4 Ip? (29-13,17,19,21), 5 pé (29-100,101,
105,106,108), 1 P,2 p,5 b Q9-1) [UCLA].
HONDURAS. Choluteca: Choluteca [elev. 0-300 ft], Dec 1945, B. Avila, 2 9 (H-9-21) [USNM];
Dec 1945, treehole, B. Avila, 2 ¢ (207E-40), 13 9 (207E-36) [UCLA]. Valle: Nacaome [el-
ev. 0-300 ft] , 20 July 1945, treehole, W. Komp, 2 9 (207E-5) [UCLA].
MEXICO. Campeche: Champoton, 10 km SW on hwy 180, elev. near sea level, 18 Aug 1966,
small treehole, height 4 ft, D. Schroeder (MEX 438), 1 Ipd (438-10), 2 Ip? (438-11,12), 1 p? (438-
100), 1 IP (438-13), 1 ?, 1 p, 6 L (438-1) [UCLA]. Nayarit: San Blas, elev. near sea level, 27 June
1956, treehole, W. McDonald (UCLA 203), 3 Ip? (203-113-115), 1 pé (203-109,110), 6 L (203-1)
[UCLA] . Oaxaca: Almoloya [elev. 800 ft], F. Knab, 22 July 1905, 1 6 (313a) [USNM].
NICARAGUA. Chinandega: Corinto [elev. near sea level] , Dec 1942, P. Woke, 1 6, 1 9 (207B-
18); 29 May 1945, well, H. Crowell, 1 6, 4 9 (H-9-24); 24 Oct 1944, steel drum, H. Crowell, 2 d,
4 2 (207B-30); 1 2, K. Maxwell [USNM] ; 19 Aug 1944, steel drum, H. Crowell, 4 6, 1 9 (207D-1)
[UCLA]. Leon: Puerto Somoza [elev. near sea level] , 13 June 1964, treehole, height 4-8 ft, A.
Quinonez (NI 4), 1 Ipd (4-113), 1 lp? (4-106), 1 pd (4-109) [UCLA]. Simonillo, nearest town
86 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
Nagarote, elev. 0-300 ft, 15 June 1964, treehole, height 5 ft, A. Quinonez (NI 11), 3 pd (11-101,
102,105), 2 p? (11-103-104); 16 June 1964, treehole, height 4 ft, A. Quinonez, 1 p? (NI 12-101)
[UCLA]. Rivas: Rivas [elev. 0-700 ft], 17 Sept 1943, treehole, P. Woke, 1 6, 2 ? (207A-22)
[USNM].
PANAMA. Panama: Bejuco, elev. 90 ft, 3 Aug 1963, old tire, height 8 ft (sic) (PA 459), 1 19
(459-101), 1 pd (459-106), 2 p? (459-107,108); 25 Aug 1963, treehole, height 8 ft (PA 534), 2
Ipd (534-109-111), 7 Ip? (534-102,104,115-119), 1 pd (534-108), 2 p? (534-105,118), 1 lp (534-
112), 1 p, 2 L, 1 1 (534-1) [UCLA]. Nueva Gorgona, elev. 60 ft, 13 Dec 1966, large treehole,
height 2 ft, O. Berlin (PA 1003), 2 pd (1003-101 ,104), 7 p? (1003-100,102,103,105,110-112),
16,3 P,1 p,21(1003-1) [UCLA].
Tehuantepec Subgroup
ADULTS. (Male known for only tehauntepec.) Head: Vertex of both sexes with
decumbent scales along longitudinal midline narrow; occiput of both sexes with e-
rect scales pale; proboscis of females longer than femur I; palpal segment 3 of male
with several long apical ventrolateral hairs forming a tuft as long as segments 4 and 5
combined; segment 4 with hairs of ventrolateral row long, closely spaced. Thorax:
Mesonotal disc of both sexes not transversely silvered; complete acrostichal and pos-
terior dorsocentral lines absent in females; acrostichal setae absent; fossal macula
of both sexes not coextensive with fossa, reduced mesally; supraalar macula not
truncate posteriorly, in females disjunct from fossal macula; ppn silver scaled; ssp
scales absent; pra hairs of females pale; upper stp and mep scale patches not con-
tiguous. Legs: Mid- and hindlegs not shaggy; mid- and hindtarsi with conspicuous
silver band; tarsus 1-I with or without narrow apical silver band; femur II with knee
spot broad, the silver scales extending basad of anterior subapical setae; tarsus 1-II
with median dark band incomplete or complete and narrow; tarsus 2-II with com-
plete apical dark band or entirely silvered; tarsus 5-II without silver scales: femur III
with basal dark band complete, narrow; tarsus 5-III without silver scales. Wing: Vein
C of females with small basal patch of silver scales, of male usually with short line
not reaching 0.5 to crossvein h; vein R and Cu of male without silver scales.
MALE GENITALIA. Sidepiece: Basal tergomesal area without dense patch of
long setae; median sternomesal area without strongly developed sclerite and with-
out tuft; specialized subapical sternal seta absent. Prosophallus: Mesal lobe with
lateral portion inclined from less than 15° to about 15° from horizontal; stem not
bowed; filament with hook not strongly angulate.
PUPA. (Known only for schroederi.) Cephalothorax: Without pale inverted V-
shaped marking; hair 5-C less than twice length of 4-C; 9-C single. Abdomen: Hair 1-
I with primary branches multiple; 2-II mesad of 3-II for 0.3 distance from 1-II
to 3-II; 3-III single. Paddle: Pigmented; apex not produced; hair 1-P shorter than
paddle.
LARVAE. Head: Hairs 5,6-C single; 7-C more than half length of 6-C; 14-C longer
than mental plate, single; bh single. Antenna: Hair 1-A single. Thorax: Hair 11-P
less than half length of 14-P; 14-P single; 3-M single; 4-M single or double; 8-T more
than half length of or subequal in length to metathoracic pleural tubercle. Abdomen:
Hair 5-I at least subequal in length to 4-I; 2-II well mesad of 4-II; 3-III single or
double; 10-III single; 3-VI single; 4-VII single; 5-VII caudad of 4-VII, more than half
length of 3-VII; 10,12-VII single. Segment VIII: Midapical comb scale with free por-
tion shorter or longer than sessile portion; hair 2-VIII single. Anal Segment: Saddle
extending around segment to nearly ventral midline, with deep ventral marginal or
Schick: Terrens Group of Aedes (Finlaya) 87
submarginal slit.
DISCUSSION. The adults of the Tehuantepec Subgroup are very similar to those
of the Podographicus Subgroup and are distinguished from the latter in the female
by the longer proboscis and broader knee spot of femur II; and in the male by the
hairier palpus and the less steeply inclined mesal lobe of the prosophallus. The lar-
vae are distinguished from the others of the Terrens Group by the more extensively
developed anal saddle and by the presence of a ventral marginal or submarginal slit.
The male genitalia and pupae do not show distinctive subgroup characters. The male
genitalia surprisingly are very similar to those of the Terrens Subgroup.
Two closely related species, tehauntepec and schroederi, comprise the subgroup.
These species are sympatric and occur on the Pacific side of the Isthmus of Tehuan-
tepec.
26. Aedes (Finlaya) tehuantepec Schick, n.sp.
Figs. 6,58,59
TYPE: Holotype é (291g), with associated pupal and larval skins and genitalia slide (671016-
60), Tehuantepec, elev. near sea level, Oaxaca, Mexico, 1 July 1905, treehole or cemented tank in
shaded part of garden at public bath house, F. Knab [USNM]. Allotype 2 (291i) with associated
pupal and larval skins, same data as holotype [USNM] . Paratypes: 5 6 (291c-f; 294j), 1 9 (291h),
same data as holotype [USNM].
Aedes (Finlaya) podographicus in part of Martini (1935:56).
Aedes podographicus in part of Howard, Dyar and Knab (1917:812-815).
Aedes insolitus in part of Dyar and Knab (1906b:189,203).
FEMALE. Head: Vertex with decumbent scales silver, narrow curved. Thorax:
(fig. 58). Legs: Femora I and II with well-developed posterior patch of silver scales
in basal half, that of femur II larger; tarsus 1-II with median dark band usually com-
plete and narrow, at most about 0.33 (incomplete in 307i); tarsus 2-II usually with
apical dark band (entirely silvered in 307i); femur III with basal dark band 0.01, sub-
apical dark band 0.33 (1 specimen); tarsus 1-III with basal silver band 0.05-0.06, a-
pical silver band 0.25-0.27 (0.38 in 307i). Wing: Vein R usually without silver scales,
sometimes with a few at base.
MALE. Head: Vertex with decumbent scales as in female; palpus about 2 label-
lum lengths shorter than proboscis. Thorax (fig. 58): Complete silver acrostichal
line present or absent. Legs: Tarsus 1-II with median dark band usually incom-
plete, when complete at most about 0.25.
MALE GENITALIA (fig. 58). Sidepiece: Length 0.34-0.37 mm. Prosophallus:
Length 0.12-0.14 mm (0.11-0.14 mm); stems essentially parallel or divergent, but
distal 0.33 in 1 specimen curved mesad (see figure); filament ratio 0.45-0.65 (0.40-
0.65). Aedeagus: Length 0.15 mm.
PUPA. Unknown.
LARVA (fig. 59). Head: Hair 7-C triple to 5-branched. Thorax: Hair 1-P double
or triple; 4,5-P double; 7-P usually triple (2-3); 4-M usually double (1-2). Abdomen:
Hair 3-III single or double. Segment VIII: Comb scales 33-54, in 4 rows, usually nar-
row spatulate, sometimes narrow ligulate; length of free portion of apical scale
0.025-0.029 mm. Siphon: Length 0.78-0.97 mm; L/S 2.38-2.63; P/L 0.49-0.59; H/L
0.55-0.67. Anal Segment: Ventral brush usually with 16 hairs (14-17); 4a-X usually
88 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
6-branched (5-7).
SYSTEMATICS. Aedes tehuantepec is distinguished from the other species of
the Tehuantepec Subgroup, schroederi, by the more numerous and more slender
comb scales of the larva which are arranged in 4 rows (usually 2 rows in schroed-
eri). The decumbent scales of the vertex of the female tehuantepec are all silver,
while in the schroederi female a submedian macula of dark scales is developed
amidst the silver scales. The constancy of this character in schroederi is open to
question since only a single female is known. The male of schroederi is unknown.
DISTRIBUTION (fig. 6). Coastal lowlands of state of Oaxaca, Mexico. Material
examined: 38 specimens; 10 6, 4 9, 2 pupae, 22 larvae; 2 individual rearings (larval).
MEXICO. Oaxaca: Almoloya [elev. 800 ft], 22 July 1905, treehole, F. Knab (308,313), 3 6
(308b), 1 ¢ (308d), 1 9 (313b) [USNM]. Salina Cruz [elev. near sea level] , 15 July 1905, treehole,
F. Knab, 1 ? (307i) [USNM]. Tehuantepec, elev. near sea level, 1 July 1905, treehole and cement-
ed tank in shaded part of garden at public bath house, F. Knab (291, type series; 294), 1
Ipd (291g), 1 Ip? (291i), 5 ¢ (291c-f; 294j), 1 2 (291h) [USNM]; 1 Sept 1965, large treehole,
height 5 ft, D. Schroeder, 6 L (MEX 333-3) — large treehole, height 8 ft, 13 L, 1 1(MEX 334-2)
[UCLA].
27. Aedes (Finlaya) schroederi Schick, n.sp.
Figs. 6,58,60
TYPE: Holotype 2? (MEX 332-10), with associated pupal and larval skins, Tehuantepec, elev.
near sea level, Oaxaca, Mexico, 1 Sept 1965, treehole, D. Schroeder [USNM].
FEMALE. Head: Vertex with decumbent scales laterad of median longitudinal
line narrow curved, dark, forming submedian macula, or silver. Thorax (fig. 58).
Legs: Femora I and II with well-developed posterior patch of silver scales in basal
half, that of femur II larger; tarsus 1-II with median dark band complete, less than
0.33; tarsus 2-II with incomplete apical dark band; femur III with basal dark band
0.05, subapical dark band 0.32; tarsus 1-III with basal silver band 0.02, apical silver
band 0.27. Wing: Vein R without silver scales.
MALE. Unknown.
PUPA (fig. 58). Paddle: Ventral midrib moderately pigmented, weakly at apex.
LARVA (fig. 60). Head: Hair 7-C double to 4-branched. Thorax: Hairs 1,4-P
double or triple; 5-P usually double (1-2); 7-P usually double (2-3); 4-M double. Ab-
domen: Hair 3-III usually single (1-2). Segment VIII: Comb scales 20-29, usually in
2 rows (2-3), stout, awl-shaped or ligulate; length of free portion of apical scale
0.024-0.032 mm. Siphon: Length 0.87-1.07 mm; L/S 2.27-2.63; P/L 0.49-0.59; H/L
0.53-0.63. Anal Segment: Ventral brush usually with 14 or 16 hairs (16-18); 4a-X 4-
6 branched.
SYSTEMATICS. See tehuantepec.
DISTRIBUTION (fig. 6). Coastal lowlands of state of Oaxaca, Mexico. Material
examined: 8 specimens; 1 2, 1 pupa, 6 larvae; 1 individual rearing (larval).
MEXICO. Oaxaca: Tehuantepec, elev. near sea level, 1 Sept 1965, large treehole, height 0.5 ft,
D. Schroeder, 1 Ip? (MEX 332-10, holotype) [USNM] — large treehole, height 5 ft, D. Schroeder,
5 L (MEX 333-4) [UCLA].
Schick: Terrens Group of Aedes (Finlaya) 89
Diazi Subgroup
ADULT. (Female only.) Head: Vertex with decumbent scales along longitudi-
nal midline narrow; occiput with erect scales pale; proboscis subequal in length to fe-
mur I. Thorax: Mesonotal disc not transversely silvered; complete acrostichal or pos-
terior dorsocentral lines absent; acrostichal setae absent; fossal macula not coex-
tensive with fossa, reduced mesally; supraalar macula not truncate posteriorly, dis-
junct from fossal macula; ppn silver scaled; ssp scales present; pra hairs pale; upper
stp and mep scale patches not contiguous. Legs: Mid- and hindlegs shaggy; mid- and
hindtarsi with conspicuous silver bands; tarsus 1-I with broad apical silver band, a-
bout 0.4; femur II with knee spot broad, the scales extending basad of anterior sub-
apical setae; tarsus 1-II] with median dark band complete, moderately broad; tarsus
2-II entirely silver scaled; tarsus 5-II without silver scales; femur III with basal dark
band complete, narrow; tarsus 5-III silver scaled. Wing: Vein C with short basal sil-
ver line, not reaching 0.5 to crossvein h.
PUPA,LARVA. Unknown.
DISCUSSION. The Diazi Subgroup is characterized by unique leg characters, the
shaggy mid- and hindlegs, the broad apical silver band of tarsus 1-I, the entirely sil-
vered tarsus 2-II, the unusually broad basal silver band of tarsus 2-III and the silvered
tarsus 5-III. Only the adult female is known.
The subgroup is monotypic and is known only from 1 locality in the state of
Veracruz, Mexico.
28. Aedes (Finlaya) diazi Schick, n.sp.
Figs. 6,56
TYPE: Holotype 2 (381-4), Cueva del Nacimiento del Agua (cave of origin of Rio Atoyac),
nearest town Penuela (elev. 3000 ft), Veracruz, Mexico, 13 July 1965, biting-landing, C.L. Hogue
[USNM]. This species is dedicated to Alfonso Diaz Najera of the Instituto de Salubridad y Enfer-
medades Tropicales, Mexico.
FEMALE. Head: Vertex with decumbent scales laterad of median longitudinal
line dark, narrow curved. Thorax (fig. 56): Mep more extensively scaled than in
other species of group. Legs: Femur I with posterior patch of silver scales poorly de-
veloped, comprising scattered scales, that of femur II well developed; tarsus 1-II with
median dark band about 0.5; femur III with basal dark band 0.07, subapical dark
band 0.37; tarsus 1-III without basal silver band, apical silver band 0.19; tarsus 2-III
with basal silver band unusually well developed, about 0.67. Wing: Vein R without
silver scales.
MALE,PUPA,LARVA. Unknown.
DISTRIBUTION (fig. 6). State of Veracruz, Mexico, at elevation of 3000 ft.
Known only by holotype female; no individual rearings.
90 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
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WN
a
Schick: Terrens Group of Aedes (Finlaya) 95
FIGURES
Distribution of the subgroups of the Terrens Group
Distribution of the species of the Thorntoni and Bertrami Subgroups
Distribution of the species of the Terrens, Alboapicus, Buenaventura and Met-
oecopus Subgroups
Distribution of the species of the Insolitus, Aitkeni and Homoeopus Subgroups
Distribution of the species of the Heteropus and Galindoi Subgroups
Distribution of the species of the Podographicus, Tehuantepec and Diazi Sub-
groups
Aedes (F.) terrens; female
Aedes (F.) terrens; male
. Aedes (F.) thorntoni; female and male mesonotum, male palpus, male genitalia
and pupa
. Aedes (F.) thorntoni; larva
. Aedes (F.) thorntoni; male genitalia and pupa. Aedes (F.) bertrami; female
mesonotum
. Aedes (F.) thorntoni; larva
. Aedes (F.) argyrothorax; female and male mesonotum, male genitalia and pupa
. Aedes (F.) argyrothorax; larva
. Aedes (F.) terrens; female and male mesonotum, male genitalia and pupa
. Aedes (F.) terrens; larva
. Aedes (F.) zavortinki; female and male mesonotum, male genitalia and pupa
. Aedes (F.) zavortinki; larva
. Aedes (F.) apollo; female mesonotum, male genitalia and pupa
. Aedes (F.) apollo; larva
. Aedes (F.) berlini; female mesonotum and pupa. Aedes (F.) braziliensis; male
mesonotum and male genitalia
. Aedes (F.) berlini; larva
. Aedes (F.) alboapicus; female and male mesonotum, male genitalia and pupa
. Aedes (F.) alboapicus; larva
. Aedes (F.) buenaventura; female and male mesonotum, female pleuron, male
genitalia and pupa
. Aedes (F.) buenaventura; larva
. Aedes (F.) metoecopus; female and male mesonotum, male genitalia and pupa
. Aedes (F.) metoecopus; larva
. Aedes (F.) insolitus; female and male mesonotum, male genitalia and pupa
. Aedes (F.) insolitus; larva
. Aedes (F.) homoeopus; female and male mesonotum, male genitalia and pupa
. Aedes (F.) homoeopus; larva
. Aedes (F.) impostor; male mesonotum, male genitalia and pupa
. Aedes (F.) impostor; larva
. Aedes (F.) gabriel; female and male mesonotum, male genitalia and pupa
. Aedes (F.) gabriel; larva
. Aedes (F.) idanus; female mesonotum, male genitalia and pupa. Aedes (F.) am-
abilis; female mesonotum
. Aedes (F.) idanus; larva
. Aedes (F.) aitkeni; female mesonotum and pupa. Aedes (F.) sumidero; female
and male mesonotum and male genitalia
. Aedes (F.) vargasi; female and male mesonotum, male genitalia and pupa
Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
. Aedes (F.) vargasi; larva
. Aedes (F.) heteropus; female and male mesonotum, male genitalia and pupa
. Aedes (F.) heteropus; larva
. Aedes (F.) galindoi; female and male mesonotum, male genitalia and pupa
. Aedes (F.) galindoi; larva
. Aedes (F.) campana; male genitalia and pupa
. Aedes (F.) campana; larva
. Aedes (F.) daryi; female and male mesonotum, male genitalia and pupa
. Aedes (F.) daryi; larva
. Aedes (F.) daryi; female and male mesonotum, male genitalia and pupa
. Aedes (F.) daryi; larva
Aedes (F.) podographicus; female and male mesonotum, male genitalia and
pupa
. Aedes (F.) podographicus; larva
Aedes (F.) podographicus; male genitalia and pupa
. Aedes (F.) podographicus; larva
. Aedes (F.) podographicus; male genitalia and pupa. Aedes (F.) diazi; female
mesonotum, tarsi 1 ,2-I-III
. Aedes (F.) podographicus; larva
. Aedes (F.) tehuantepec; female and male mesonotum and male genitalia. Aedes
(F.) schroederi; female mesonotum and pupa
. Aedes (F.) tehuantepec; larva
Aedes (F_) schroederi; larva
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Schick: Terrens Group of Aedes (Finlaya) 157
INDEX TO SCIENTIFIC NAMES
Aedes Meigen, 3, 5
aitkeni Schick, 7, 10, 21, 25k, 27k, 61-62; 4
39
Aitkeni Subgroup, 18, 20, 25k, 61; 1, 4
alboapicus Schick, 5, 6, 8, 11, 17, 22, 24k,
25k, 28k, 30k, 50-51; 3, 23, 24
Alboapicus Subgroup, 17,18, 19,20, 25k, 28k,
49-50,52;1, 3
amabilis Schick, 10, 22, 27k, 30k, 68-69; 5,
a7
apollo Schick, 20, 22, 26k, 28k, 31k, 40, 42,
44, 46-48; 3, 19, 20
argyrothorax Bonne-Wepster & Bonne, 3,5, 6,
1, 2%. £2, 13; LOTT Te 21 22.2 4 2k,
28k, 31k, 33, 36-38, 40; 2, 13, 14
argyrothorax of authors, 63
berlini Schick, 22, 25k, 27k, 31k, 40, 48-49;
Sat 22
bertrami Schick, 22, 25k, 27k, 30k, 38-39; 2,
il
Bertrami Subgroup, 17, 18, 19, 25k, 38; J, 2
braziliensis Gordon & Evans, 3, 6,7, 22, 24k,
29k, 30k, 40, 43-44; 3, 2]
buenaventura Schick, 7, 10, 13, 16, 17, 25k,
20k, 30k, 52-53: 3, 25, 26
Buenaventura Subgroup, 17, 18, 19, 20, 25k,
28k, 51-52, 54; 1, 3
eampana Schick, 9, 10,12, 15,21, 22, 27k,
29k, 31k, 77, 78, 79, 80;5, 46, 47
conservator, 21
Culex of, 5
daryi Schick, 15,26k, 29k, 30k, 77, 78, 80-81;
5, 48,49, 50, 51
diazi Schick, 10, 11, 17, 26k, 27k, 89; 6, 56
Diazi Subgroup, 18, 19, 26k, 89; J, 6
Finlaya Theobald, 3,5, 16, 24
gabriel Schick, 7, 13, 22, 27k, 29k, 31k, 68,
69-70, 73; 81; 5,35, 36
galindoi Schick, 5, 6,9, 10, 12, 15, 25k, 27k,
29k, 30k, 77-79, 80; 5, 44, 45
Galindoi Subgroup, 18, 19, 20, 21, 26k, 29k,
76-77; 1, 5
Geniculatus-Group, 17
Gualteria Lutz, 5
Haemagogus of, 5
heteropus Dyar, 3, 5, 6, 20, 27k, 30k, 65, 68,
73, 74-76;5, 42, 43
Heteropus Subgroup, 18,19, 21, 26k, 29k, 54,
63, 67-68, 82; 1, 5
homoeopus Dyar, 3, 6, 7, 8, 12, 13, 22, 27k,
28k, 31k, 63-66, 67, 75, 76; 4, 31, 32
Homoeopus Subgroup, 12, 18,19, 20, 21, 26k,
27k, 54, 57, 62-63, 68, 82; 1, 4
idanus Schick, 22, 27k, 29k, 30k, 68, 70-72,
TI DINSLNSO.
impostor Schick, 8, 24k, 28k, 31k, 63, 65, 66-
67; 4, 33, 34
insolitus (Coquillett), 3, 5,6,8,9,11, 15, 22,
24k, 26k, 28k, 31k, 57-61; 4, 29, 30
insolitus of authors, 33, 45, 83, 87
Insolitus Subgroup, 17, 18, 19, 20, 25k, 28k,
56-57; 1, 4
kompi Vargas & Downs, 16
laternarius (Coquillett), 5, 6,57
Longipalpis-Group, 17
Lophoceraomyia, 15
mediovittatus (Coquillett), 5
metoecopus Dyar, 3, 6, 7, 11, 22, 25k, 29k,
31k, 54-56; 3, 27, 28
Metoecopus Subgroup, 17, 18, 19, 20, 25k,
29k, 53-54, 63, 68, 82; 1, 3
oswaldi (Lutz), 5, 6, 41
oswaldi of authors, 5, 36, 44, 57, 64
podographicus Dyar & Knab, 3, 5, 6, 10, 20,
21 26k, 29k. 31k: 82-8676, 52, 22, 04,20,
D657 |
podographicus of authors, 6, 41, 46, 55, 57,
64, 74, 87
podographicus complex, 11, 84; 6
Podographicus Subgroup, 13, 18, 19, 20, 21,
26k, 29k, 54, 63, 68, 81-82, 87; 1, 6
poicilius (Theobald), 5
schroederi Schick, 15, 16, 26k, 27k, 31k, 86,
87, 88-89; 6, 58, 60
scutellalbum, 16
158 Contrib. Amer. Ent. Inst., vol. 5, no. 3, 1970
sumidero Schick, 8, 22, 27k, 30k, 68, 72-73;
5, oY
tehuantepec Schick, 15, 16, 26k, 29k, 31k,
86, 87-88; 6, 58, 59
Tehuantepec Subgroup, 16, 18, 19, 26k, 29k,
86-87; 1, 6 |
terrens (Walker), 3, 4,5, 6, 7,8, 10, 16, 17,
21, 22, 23, 26k, 28k, 31k, 40, 41-43, 47,
48; 3, 7, 8, 15, 16
terrens of authors, 33, 43, 46,55, 57, 64, 74,
79, 83 |
Terrens Group, 3,5,9, 15, 16, 17, 18, 19, 20,
21, 23-24, 32, 33, 34, 42, 49, 57, 61, 72,
81, 87
Terrens-Group: Gualteria, 5, 16
Terrens-Gubernatoris-Group: Gualteria, 5
Terrens Subgroup, 12, 17, 18, 19, 20, 21, 25k,
28k, 33, 37, 39-40, 42, 50, 52,57, 87; 1, 3
thorntoni Dyar & Knab, 4, 5, 6, 8, 16, 17,
18, 21, 25k, 28k, 31k, 33-36; 2, 9, 10, 11,
£2
thorntoni of authors, 45
Thorntoni Subgroup, 10, 17, 18, 19, 20, 21,
25k, 28k, 32-33, 38, 40; 1, 2
triseriatus (Say), 5
vargasi Schick, 10, 22, 27k, 30k, 31k, 68, 73-
74, 84; 5, 40, 41
zavortinki Schick, 5, 6, 11, 12, 20, 21, 26k,
29k, 31k, 40, 44, 45-46, 47; 3, 17, 18
a
No.
(Continued from inside front cover)
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Contributions
of the
American Entomological Institute
Volume 5, Number 4, 1970
CONTRIBUTIONS TO THE MOSQUITO FAUNA OF
SOUTHEAST ASIA, - V
Genus Aedes, subgenus Diceromyia Theobald in
Southeast Asia.
by
John F. Reinert
MAY 1 8 1970
LiBRARIE?
CONTRIBUTIONS
of the
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Contributions
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American Entomological Institute
Volume 5, Number 4, 1970
CONTRIBUTIONS TO THE MOSQUITO FAUNA OF
SOUTHEAST ASIA. - V
Genus Aedes, subgenus Diceromyia Theobald in
Southeast Asia.
by
John F, Reinert
CONTENTS
PCE ROO UG TION ie ee ee a ee ee
Genus Aedes, Subgenus Diceromyia .....+-e.e«ee+seeces
LIST OF THE SPECIES IN SOUTHEAST ASIA .......
KEYS TO THE ORIENTAL SPECIES . 45.6.2 06 3 2 e's
MALES AND FPREMALIS . yee we ee ee
ee a ee a ee ee ee ee
De ke ge ie ae eh Reese ea a wel
DESCRIPTIONS OF THE SPECIES OCCURRING IN
OU Pelee l eA oo a ee ee ee ee ew,
FIORELSCOL MARGDELY 2 in Riper do 6 ce ere ee ey
ene BOWS Fs OE PF eee eS Fe es
Diaiylepidus Knight and: Aull... sk ee ee ee
pepcripes HOwards oc We no a ee ws
Per eee Oe ei ke Sc ek Se RS ee
OO ea fe see Ce ee ok ese bo ae ee
SOIT Techy ee ee ee a ea ew
BCR ae I ies ee ae as ek ee le ee
PT A a es a be ee Re es nee ake
APPENDICES
TABLE 1. Present knowledge of the Diceromyia species
in the Ethiopian Zoogeographical Region ......
TABLE 2. Present knowledge of the Diceromyia species
INDEX
in the Oriental Zoogeographical Region. ......
CO ©O ITO OO he
CONTRIBUTIONS TO THE MOSQUITO FAUNA OF SOUTHEAST ASIA. V.
GENUS AEDES, SUBGENUS DICEROMYIA THEOBALD
IN SOUTHEAST ASIA. !
By
John F. Reinert
INTRODUCTION
The subgenus Diceromyia was originally described by Theobald
(1911: 151) as a distinct genus based on a single African species. Several
Oriental species of the present subgenus were initially included in Aedes
(Skusea) Theobald by Edwards (1922: 272) but later (1929: 341) he placed them
in Dendroskusea, as a separate subgenus of Aedes. Edwards (1932: 172)
combined the existing 4 African and 5 Oriental species under the genus Aedes
subgenus Diceromyia and separated the species into group A (African species)
and group B (Oriental species). At the present time 8 species from the Ethi-
opian Zoogeographical Region (Appendix: Table 1) and 11 species from the
Oriental Zoogeographical Region (Appendix: Table 2) are recognized as be-
longing to Aedes (Diceromyia), For a discussion of the inclusion of kanarensis
see Mattingly (1965:66).
The present paper deals with the 7 species from Southeast Asia of
which one is described as new and the male of platylepidus Knight and Hull
and the female of whartoni Mattingly are described for the first time. Keys
to the adults, pupae and larvae of the Oriental species are given. For the de-
tailed descriptions of the Oriental species not found in the Southeast Asia area
see: Khokhar and Tariq (1966: oe Reuben (1967: 234) vama-
chandvai; Barraud (1934: 275, 444) micropterus and kanarensis. Descriptions
and a key to the adults of most of the African species are given by Edwards
(1941: 214). He also gives the descriptions to 3 of the pupae. A key to pupae
of the African species is given by De Meillon, Parent and Black (1945: 93).
The larvae of 6 species from Africa are described and illustrated by Hopkins
(1952: 116, 213) and included in a key to the Aedes. Other important papers
dealing with African species of Diceromyia not mentioned above are: Doucet
(1951: 69) gvassei; Wolfs (1958: 298) bananea; and De Meillon (1943: 94) zethus.
Abbreviations used in the references to literature conform to the
World List of Scientific Periodicals, 3rd ed., Academic Press, 1952, An
asterisk following the abbreviations used (? = female, “ = male, P = pupa, L
= larva) indicates that at least some portion of that sex or stage is figured.
Distribution records are indicated as follows: countries are in capital letters,
I this work was supported by Research Contract No. DA-49-193-MD- 2672
from the U.S. Army Medical Research and Development Command, Office of
the Surgeon General and carried out at the Southeast Asia Mosquito Project,
Washington, D.C.
2Major, MSC, U.S. Army, Department of Entomology, Walter Reed Army
Institute of Research, Walter Reed Army Medical Center, Washington, D.C.
20012.
2 Contrib. Amer. Ent. Inst., vol. 5, no. 4, 1970
where known administrative divisions are in italics and place names have the
first letter capitalized.
Specimens of the following species of Aedes (Diceromyia) have been
examined by me: adersi (Edwards), fascipalpis (Edwards), franciscoi
Mattingly, furcifer (Edwards), ivengari Edwards, micropterus (Giles), peri-
skelatus (Giles), platylepidus Knight and Hull, ramachandrai Reuben, reginae
Edwards, scanionin.sp., taylori Edwards, whartoni Mattingly, and zethus
De Meillon and Lavoipierre.
GENUS AEDES MEIGEN
SUBGENUS DICEROMYIA THEOBALD
Diceromyia Theobald 1911, Rep. Wellcome trop. Res. Lab. 4(B): 151. Type
saber ing re eign africana Theobald; Edwards 1917, Bull. ent.
Res. 7: 214.
Aedes Skusea) Edwards 1922, Indian J. med. Res. 10(1): 272 (partim);
Barraud 1928, Indian J. med. Res. 16: 357 (partim).
Aedes (Dendroskusea) Edwards 1929, Bull. ent. Res. 20: 341. Type species:
Culex micropterus Giles.
Aedes (Diceromyia) Edwards 1932, Genera Insectorum, Fasc. 194: 172;
Barraud 1934, Fauna Brit. India, Diptera 5: 271; Edwards 1941,
Mosq. Ethiopian Region 3: 214; Hopkins, 1952, Mosq. Ethiopian
Region 1: 115; Mattingly 1959, Culic. Mosq. Indomalayan Area 4: 34.
FEMALE. Head. Torus with patch of fine short hairs, scales or both
mesally; Ist flagellomere usually with a few small scales; palpus 0. 2 -0.3
length of proboscis; proboscis slender, from slightly longer to slightly shorter
than fore femur; eyes usually contiguous; orbital and frontal bristles well devel-
oped; orbital line covered with pale scales; decumbent scales of vertex mainly,
to all, broad, forming pale and dark patches, erect scales few to numerous (ab-
sent in platylepidus); erect scales always numerous on occiput. Thorax.
Scales of scutum broad, narrow or both; scutellum with broad flat scales (ex-
cept in flavicollis which has narrow scales, furcifer and faylori, which have
broad scales on the lateral lobes and broad with a few narrow ones on the
center lobe); anterior promontory, humeral and supraalar bristles always
present, other bristles on scutum well developed to absent, scutellum with
well developed bristles; postnotum bare; anterior pronotal lobes widely sepa-
rated with well developed bristles and scales; pleuron with a patch of broad
white scales on each of the following areas: subspiracular (except in platy-
lepidus), propleural, sternopleural, and mesepimeral; all species,except
platylepidus, have broad or narrow pale scales on the paratergite; bristles
present on the following pleural areas: posterior pronotal, propleural, post-
Spiracular, upper and lower sternopleural, prealar, and upper mesepimeral
(lower present or absent); upper edge of meron above base of hind coxa. Legs.
Fore and mid coxae with pale scale patches; femora scaling varied, fore and
mid femora swollen in comparison to hind femur, tibial scaling varied; tarsi
with tarsomeres dark, spotted or basally banded; fore and mid tarsal claws
toothed (African species) or simple (Oriental species), hind tarsal claws
simple. Wing. Usually dark and pale scaled, pale scales may be restricted
to a basal patch on the costa (scales all dark on wings of micropterus and
reginae); dorsal scales broad at least on costa, subcosta and usually a portion
of the radius and cubitus; cells Ro and Mg longer than their stems; anal vein
terminating well beyond fork of cubitus; alula with scales along margin anda
Reinert: Aedes (Diceromyia) in Southeast Asia 3
small apical patch in some species; squama with hair fringe; 1-3 remigial
bristles present. Halter. With pale stem and dark scaled knob (some species
with a few pale scales at base of knob). Abdomen. Terga scaling variable
but with some pale scales, bands or patches present; tergum I with a rectan-
gular patch of white scales on the lateral tergite; sterna pale scaled or with
pale basal bands; segment VII extended or retracted into segment VII, when
segment VIII is extended the cerci are retracted within it and not dorsally
visible. Terminalia, Cerci short and broad; spermatheca triple.
MALE. Similar to female in general habitus. Head. Antenna plu-
mose with hairs directed mainly dorsally and ventrally; palpus with apical and
subapical segments very short, slightly thickened, and usually turned some-
what downward; 3 distal segments with a few hairs; palpus slightly longer to
slightly shorter than proboscis (0.6 length of proboscis in platylepidus). Legs.
Fore and mid tarsal claws unequal with larger claw toothed, hind claws smal-
ler, equal and simple. Terminalia. IXth tergum bilobed with several dorsal
hairs on each lobe; basimere usually broad (may be moderately broad in some
species) with several apical long bristles and numerous small bristles on dor-
sal and ventral sides (few smaller bristles in fascipalpis); distimere with
spiniform attached subapically (except in micropterus); basal mesal lobe with
fine short hairs or with moderately to well developed bristles, connected
sternally by a narrow membranous Strip to its mate; phallosome with aedeagus
divided into two lateral plates each bearing several teeth (number of lateral
teeth reduced in flavicollis and grassei); paraproct simple, with apex, lateral
margin and base moderately sclerotized, cercal setae absent.
PUPA. The pupae of the Oriental species do not, at this time, pre-
sent any clear-cut subgeneric characters. They do, however, have certain
Similarities. Cephalothorax, Hairs 6, 8, 9, 11-C single; trumpet short
(moderately long in whartoni). Abdomen. Hair 9-I-VI is small and stout
while 9-VII is stout and at least one-half the length of segment VII; 9-VIII is
stout and longer than segment VI; hairs 2, 3-I-VIL are single; most of the
other abdominal hairs are usually forked or branched and extremely variable;
paddles oval with hair 1-P well developed.
LARVA. Head, Hair 1-C long, slender and pointed; 3-C single; 4,
7-C multiple; 6-C with 1-3 branches; 6-C anterior to 5-C and on a more-or-
less longitudinal line with it; 4-C anteromesal to 5-C and usually posteromesal
to 6-C; antenna short (moderately long in whartoni), smooth or spiculate and
hair 1-A single to 5-branched; mouthbrushes in some species with pectinate
setae; mentum with 8-12 teeth on each side. Thorax. Meso- and metapleural
tubercles with variously developed spines. Abdomen. Comb consisting of 4-
25 scales arranged in 1 or 2 rows, scales usually with a fine lateral fringe;
siphon short (moderately long in whartoni), hair 1-S single to 3-branched and
attached beyond apical pecten tooth at about the middle of the siphon; pecten
with 5-14 evenly spaced teeth, each with denticles on ventral margin to apex;
hair 1-X single to 5-branched; 2-X with 2-7 branches; 3-X single; 8-11 hair
tufts in ventral brush, with usually 1-3 tufts proximad of grid; saddle incom-
plete, usually covered with fine spicules; anal papillae with rounded apices
and varying in length.
Oriental species also have the following additional characters in com-
mon. Head, 5-C long and single (sometimes double in peviskelatus); 8-C
single; 9-C single (single or double in periskelatus); 10-C usually double; 11-C
with 5-18 branches; 12-C with 2-4 branches; basal maxillary hair stout and
pare (3-5 branched in periskelatus); antenna with hair 5-A rectangular and
eaflike.
4 Contrib. Amer. Ent. Inst., vol. 5, no. 4, 1970
DISTRIBUTION. ! Species of the subgenus Diceromyia are about
evenly divided between the Ethiopian Zoogeographical Region (8 species) and
the Oriental Zoogeographical Region (11 species). In the Ethiopian Region
species have been collected from: BOTSWANA (Bechuanaland), DEMOCRAT-
IC REPUBLIC OF THE CONGO (Belgian Congo), ETHIOPIA, GAMBIA, GHANA
(Gold Coast), KENYA, MALAGASY REPUBLIC (Madagascar), MALAWI (Nya-
saland), NIGERIA, PORTUGUESE GUINEA, RHODESIA (Southern Rhodesia),
SENEGAL, SOUTH AFRICA, SUDAN, TANZANIA (Tanganyika), UGANDA,
UPPER VOLTA, ZAMBIA (Northern Rhodesia) and from the Oriental Region:
BURMA, CEYLON, INDIA (mainland and Nicobar Islands), INDONESIA, WEST
MALAYSIA, PHILIPPINES, SINGAPORE, and THAILAND.
ZOOGEOGRAPHY. The geographical distribution of the subgenus
Diceromyia is confined chiefly to the savanna and forest fringe areas of the
Ethiopian and Oriental Zoogeographical Regions. Eight species are limited
to the former area and 11 to the latter with no exchange of species between the
two regions. The subgenus may have evolved in the Indian area and then dis-
persed to the Ethiopian Region when the climate and vegetation of the inter-
vening area were more propitious. As the climate changed, the favorable en-
vironmental corridor ceased to exist and the populations of the two areas dis-
persed and evolved in their respective regions. Their present ranges corre-
spond more or less with definite climatic and vegetation zones found in these
regions,
A more detailed discussion on the zoogeography of the subgenus is in
press.
TAXONOMIC DISCUSSION. The subgenus Diceromyia, although be-
ing quite varied in chaetotaxy and pilotaxy, possesses a combination of charac-
ters that allows it to be separated from the other subgenera of Aedes. The
male exhibits the most distinctive features, these being: palpus with the two
terminal segments very short; antenna with the plume hairs directed mainly
dorsally and ventrally; the terminalia which have the aedeagus divided into
two lateral plates each bearing several teeth and the absence of cercal setae
on the paraproct (the characters of the terminalia fit into Section B, Subsec-
tion 3 of the classification system given by Belkin (1962: 326). A similar pal-
pus is found in some Stegomyia, Ochlerotatus and Finlaya while the antenna is
typical of Aedimorphus. The toothed aedeagus and the paraproct are similar
to those of Aedimorphus and Stegomyia.
Other characters of the adults that are constant are the presence of
pale scales on the paratergite and the vertex with two or three dark areas al-
ternating with pale patches (except in platylepidus which has the paratergite
bare and different markings on the vertex). These features are, however,
found in many species of Stegomyia and Aedimorphus. A number of species
of Diceromyia from both the African and Oriental regions have the wing scales
very broad and the leg markings similar to species of Mansonia,
The females have the cerci short and broad and the spermatheca tri-
ple. Females of the Oriental species have the tarsal claws simple while those
of the African species are toothed. Female specimens can usually be sepa~
rated from those of the other subgenera of Aedes by a combination of the char-
acters mentioned.
The pupae of the Oriental species are very similar, especially in the
chaetotaxy, shape of the trumpet and the structure of the paddle. However,
1 Former name of country enclosed in parenthesis.
Reinert: Aedes (Diceromyia) in Southeast Asia 5)
whartoni is distinctive and differs in having a moderately long trumpet, paddle
with tiny spicules on the proximal 0.5 of the outer margin, and in the struc-
ture of hairs 3-C, 9-VIII and 1-P.
The larval characters listed for the subgenus are shared by members
of Stegomyia and Aedimorphus. Larvae of Diceromyia can usually be sepa-
rated from these two subgenera by the shape of the comb scales and pecten
teeth or a combination of the other characters given for the subgenus.
BIOLOGY AND MEDICAL IMPORTANCE, Two species of African
Diceromyia have been shown to be potential vectors of human pathogens. A
number of other African and Oriental species are known to feed on man but
have not been investigated for pathogen transmission. Lewis (1943: 73) lists
taylovias an important potential vector of yellow fever virus in the Sudan and
also considers furcifer as being suspected in this connection. He mentions
that these two species were collected in large numbers at ground-level biting
man and that they are probably very active fliers or long-lived. Mattingly
(1949a, b) Haddow et al. (1951) and Van Someren et al. (1955), however, re-
cord these species feeding in or below the canopy. Haddow (1961: 324) states
that the furcifer-taylori group is arboreal and crepuscular but that these mos-
quitoes will also bite freely at ground-level when forest or brush is scarce or
absent and that they are very resistant to drought and heat. He also mentions
that they rarely enter huts and tents even in areas where they are very pre-
valent outside. Haddow records the biting habits of another species, adersz,
as nocturnal and arboreal. The interesting mating behavior offaylovi is re-
corded by Hamon et al, (1955). These researchers observed males of this
species mating with females while the latter were actually engorging.
During disease transmission experiments in Southern Rhodesia,
Paterson and McIntosh (1964: 54), used wild caught females of the furcifer-
taylori group and showed that they were efficient vectors of chikungunya virus
between monkey and monkey and between monkey and mouse. They also re-
ported the accidental transmission of chikungunya virus to two workers in the
laboratory by members of the furcifer-iaylori group.
The adults of scanloni n. sp. in Thailand were collected both in light
traps and resting on the inside of houses. A number of investigators report
species of Oriental Diceromyia feeding on man. Scanlon and Esah (1965: 143)
record iyengavi biting man in Thailand in the forest fringe and scrub areas.
Specimens of vamanchandrai were collected during all seasons of the year in
India either in vegetation or biting man according to Reuben (1967: 236).
Mattingly (1959: 43) states the holotype female of fvanciscoi was collected in a
trap using human bait.
Hopkins (1952) summarizes the breeding habitats of African species
as primarily tree holes but specimens were also collected in rot holes, axil
of a banana leaf, water pot and an iron tank. A. furcifey and faylori were re-
corded breeding in rock pools by Lewis (1943: 69), while Van Someren et al.
(1955: 482) collected taylori larvae from pineapple tops and steps cut into coco-
nut palm trees and adersi larvae were taken from bamboo pots, coconut shells
and domestic utensils. The Oriental species are primarily bamboo and tree
hole breeders as reported by Barraud (1934). Knight and Hull (1951: 201) also
collected larvae of platylepidus from a fallen coconut spathe and a log depres-
sion.
The eggs of Diceromyia are drought-resistant. This adaptation en-
ables the subgenus to survive during dry periods. In Africa eggs of furcifer
that had been laid in bamboo pots more than 14 months earlier were flooded
and hatched, demonstrating that the eggs of this species are very resistant to
desiccation (Muspratt, 1955: 174). Mattingly (1959: 2, 43) reports that in
Contrib. Amer. Ent. Inst., vol. 5, no. 4, 1970
Malaya eggs of franciscot were laid in bamboo pots hung at about 30 feet from
the ground in a mango tree.
5(1).
6(5).
7(5).
8(7).
LIST OF SPECIES IN SOUTHEAST ASIA
. Aedes (Diceromyia) franciscoi Mattingly
Aedes (Diceromyia) iyengavi Edwards
Aedes (Diceromyia) platylepidus Knight and Hull
Aedes (Diceromyia) punctipes Edwards
Aedes (Diceromyia) reginae Edwards
Aedes (Diceromyia) scanloni n. Sp.
Aedes (Diceromyia) whartoni Mattingly
ID OMB Oo bor
KEYS TO THE ORIENTAL SPECIES!
MALES AND FEMALES
Presculeliar DFisclcs BHOSCHL .. oe. «. andi.) a0 68% ec bee © e
Prescure lar DristicS OrGGent ai... 6. 6 ease ue e: mce jane bucene
Anterior mesonotum covered with broad scales ....cecee
Anterior mesonotum covered with narrow scales....s.e-ev.«e
> Oo O1 po |
Paratergite and posterior pronotum bare .... platylepidus (p.12)
Paratergite and posterior pronotum with broad
scales @ @ e @ @ @ e @ @ @ @ @ @ e @ e e @ e @ tC) franciscot (Gon 8)
First hind tarsomere with a basal white band, rest
of hind tarsomeres dark ... scipics eibatedi ake Te kanarensis
Hind tarsomeres 1, 2, 3 and 4 with basal white |
Oe a ek oo sw ee oe whartoni (p19)
Scales on wing all dark @ @ e @ e @ e@ @ e@ @ @ @ € @ e ® @ @ ® @ @ 6
Ae ON WI A gg ose) bes) eet Soe w vehi eee cece bk % 7
Pleural integument dark brown; apices of all
femora with lateral white spots; lateral surface
of head dark scaled ...... ‘ « ete «.,,. MIiCcyoplerus
Pleural integument pale yellow; apices of all
femora dark scaled; lateral surface of head
Cee reginae (p.15)
Tarsi, palpus and proboscis each without a
dorsal line of white scales. ... otigl o wlna ae bes 8
Tarsi, palpus and proboscis each with a dorsal
TUS Gl WO SCOICS 6 ee he seen’ ecb tie, o08 rama chandrai
Wing with pale scales on bases of costa and at
TRO SULON ONG OLIEE VOU ons ice, bcm cael ei loie Wie 0 eee a
Wing with pale scales only on base of costa... iyengari (p. to
t The keys include Oriental species which may yet be found to occur in
Southeast Asia.
Reinert: Aedes (Diceromyia) in Southeast Asia 7
9(8).. Wing with pale scales on base of cubitus. .......2 cece 10
Wing with pale scales on basal third of subcosta,
scales on cubitusdark 209 Ge Serre ee periskelatus
10(9). Wing with pale scales on base of radius; palpus
with apical pale band’s -.°. 2°. i.e 3. st 6 SeQnionz Nesp. (p. 7"
Wing with dark scales on radius; palpus dark .. punctipes (p.14
PUPAE
1. Paddie liair PP Ginvle i"... oe a se ee a ee
Paddle hair 1-P with 2-5 branches ....... whartoni (p. 20)
2(1). Paddle edge with fine hairs; hair 1-II double ......... 3
Paddle edge with spicules; hair 1-1 SUNOIe eee 4
micropterus;, reginae (p.16)
3(2). Hair 5-VI, equal to or longer than length of
segment Vil: 9-VHI Single’ or double .°3. 06. 3. Se se 4
Hair 5-VI shorter than length of segment VII;
9-VIII with three branches ........e2+..e /fvanciscoi (p. 9 )
4(3). Hair 1-V single or double; 9- VII one-half
leneth of Segment VIP 6 ee re eS ee iyengari (p.11)
Hair 1-V with 3-6 branches; 9-VII three-
fourths length of segment VII .......e.-. periskelatus
LARVAE
de Antenna with spicules on basal one-half, hair
1=A doubles... ee@eee#e# e@# @ @® @® @ @ 2
Antenna without spicules, hair 1-A single Peed 6a ea 64 3
2(1). Basal maxillary hair single; saddle hair 1-X
Single ., ey Ae eo whartoni (p. 20)
Basal. maxillary hair with 3-5 branches:
saddle hair 1-X with 3-4 branches ...... periskelatus
3(1). Siphon hair 1-S single; saddle hair 1-X
single or double ». . ete ee era ee a 4
Siphon hair 1-S with ee branches; “saddle
hair 1-X with 3-5 branches . ... micropterus, reginae (p.16)
4(3). Comb composed of 5-10 scales arranged in
one row.) ss. a ei iyengari (p.12)
Comb composed. of 15- 25 ‘scales arranged
in two rows ora SO ee ee 28 ea ee enc isear Ce, 2)
Immatures of kanarensis, punctipes, vamachandyvai and scanloni n. sp. are
unknown. The larva of platylepidus is not known with certainty.
8 Contrib. Amer. Ent. Inst., vol. 5, no. 4, 1970
DESCRIPTIONS OF THE SPECIES OCCURRING IN SOUTHEAST ASIA
AEDES (DICEROMYIA) FRANCISCOI MATTINGLY
Figs. 1, *9; 7, & terminalia; 10, 11, pupa; 14, larva.
Aedes (Diceromyia) franciscoi Mattingly 1959, Culic. Mosq. Indomalayan
Area 4: 42 ms , 9*, L, P); Mattingly 1961, Culic. Mosq. Indomalayan
Area 5: 59 (c'*, P, iL),
FEMALE, Head, Antenna dark brown, longer than the proboscis,
torus with a patch of pale scales and short fine hairs mesally, flagellomere 1
with a few small pale scales and equal in length to flagellomere 2; clypeus
dark, bare; palpus dark with apex pale, 0. 25 length of proboscis; proboscis
dark with a few pale scales ventrally at about middle, slightly longer than fore
femur; vertex with alternating dark and pale broad decumbent scale patches,
two patches of pale scales separated by a central dark area, another pale patch
on each side with dark scales below, a few dark erect scales posterior to or-
bital line and numerous short ones on occiput. Thorax. Anterior mesonotum
covered with loosely attached broad brown scales with a few paler ones inter-
mixed, broad white scales extending around anterior margin to about scutal
angle; posterior mesonotum with overlapping longer dark brown scales on
posterior portion extending laterally and anteriorly over supraalar bristles,
same type of scales extending over and completely covering scutellum, a few
broad white scales above paratergite; anterior promontory, humeral, supra-
alar and scutellar bristles well developed, others absent; pleural integument
dark brown; anterior pronotal lobe covered with bristles and long broad white
scales; upper posterior pronotum with broad brown scales, middle with broad
white scales and 4 posterior bristles; propleuron with broad white loose fitting
scales, and 5-6 bristles; prosternum bare; postcoxal membrane with broad
white scales on upper 0.5; spiracular area bare; postspiracular area with 4
bristles; subspiracular and hypostigial areas with broad white scales; para-
tergite with broad white scales along margin; sternopleuron with upper and
lower broad white scale patches and upper and posterior bristles; prealar area
with an upper patch of bristles and broad white scales; mesepimeron with a
patch of broad white scales over anterior, upper and posterior areas, a patch
of upper and 2-4 middle bristles. Legs. Coxae with white scale patches, fore
and mid coxae also with some dark scales; trochanters white scaled; femora
with two more or less distinct subapical white bands anteriorly, other white
scale patches present; fore and mid tibiae with 5 lateral white spots anda
small apical white spot, hind tibia with subapical white band, a dorsoapical
and 3 lateral white spots; fore and mid tarsi with tarsomere 1 having a narrow
basal white band, a small dorsal white spot at about the middle and one at apex,
tarsomeres 2-5 with small dorsobasal pale spots (spots indistinct on 4 and 5
in some specimens); hind tarsomere 1 with a dorsoapical pale spot, 3 white
bands (one at base, one at about 0.3 of the length and one at about 0.6 of the
length), tarsomeres 2 and 3 each with a narrow white basal band, a dorsoapi-
cal spot and a tiny dorsal white spot at about the middle, tarsomeres 4 and 5
each with a dorsobasal pale spot. Wing. Dorsal veins covered with broad
brown scales and with a patch of white scales on anterior margin of costa
from the base to the humeral crossvein and a few at base of the remigium;
broad brown scales along posterior margin of wing above fringe scales; alula
with broad brown scales on fringe and a small dorsoapical patch; 1-2 remi-
gial bristles. Halter. With stem pale and knob brown scaled, a few pale
scales ventrally. Abdomen. Terga with brown and pale markings, tergum I
with a large pale scale patch covering most of dorsal surface; terga II-VI each
Reinert: Aedes (Diceromyia) in Southeast Asia 9
with a pair of dorsolateral semicircular pale bands with dorsal ends touching
in middle on terga II-IV (bands more distinct on anterior terga), terga II-VI
each with a narrow dorsoapical fringe of broad yellow scales (yellow scales
more numerous on posterior terga), tergum VII with a few scattered pale
scales; posterior margins of terga and sterna fringed with golden hairs (hairs
more numerous on sterna); segment VIII completely retracted into segment
VII.
MALE. Similar to female in general habitus. Head. Palpus with
basal pale band on terminal segment, a few pale scales at about the middle of
segments 1-3; palpus approximately equal to proboscis; proboscis with a nar-
row pale band at about the middle. Wing. Pale scaling on wing somewhat re-
duced. Abdomen. Tergal scale patterns more distinct than on female; ter-
gum VIII with dorsobasal white scale patch. Terminalia, Tergum IX bilobed
with 13-15 bristles on each lobe, the outer 7-8 bristles longer; basimere short
and broad with a patch of moderately long flattened bristles on the tergomesal
margin extending from below the apex to the base, a small flattened lobe with
fine hairs extending above these bristles to apex, numerous long bristles on
dorsal surface and shorter ones on ventral surface, scales numerous on later-
al margin and extending onto dorsal and ventral surfaces; distimere shorter
than basimere by about length of the spiniform, thickened and tapering to a
blunt apex, spiniform dark, attached subapically and extending slightly beyond
tip of distimere; basal mesal lobe with 15-18 moderately long flattened bris-
tles, basal connecting fold covered with short hairlike spicules; phallosome
with aedeagus divided into two lateral plates each with 7-9 teeth; sternum IX
with 4-5 bristles along posterior margin.
PUPA. Cephalothorax. Hairs 1-3, 10, 12-C single; 4, 5, 7-C single
or double. Abdomen, Hair 1-II-III forked or double; 3-III single, shorter than
the length of segment III; 1-V double or forked; 5-VI single, shorter than the
length of segment VII; 9- VI one-half to two-thirds length of segment VII; 9-
VIll with 3 branches, from slightly shorter to slightly longer than the length of
segment VII. Paddle, With hairlike spicules along most of the outer (distal
0. 75) and inner (distal 0.5) margins; paddle hair 1-P single.
LARVA. Head, Hair 4-C with 14-25 branches; 6-C double; 7-C with
6-10 branches; 10-C single with 8-13 branches; 12-C with 2-4 branches; an-
tenna short, without spicules and with hair 1-A single and attached at about
0.62 distance from base. Thorax. Hair 9-M stellate with 6-7 branches; hair
4-T with 2-3 branches and hair 5-T single or double. Abdomen, Hair 2-I-
VIII single; hair 1- VII with 2-3 branches; comb consisting of 15-25 scales in
two irregular rows, most of the scales in anterior row smaller, scales witha
fine fringe along margins and the bluntly pointed apex; siphon index approxi-
mately 2.0-2. 2; hair 1-S single and reaching apex of siphon (minus valves);
pecten with 9-16 teeth, with denticles along ventral and apical margins; hair
1-X single or double and about 1.0-1.5 length of saddle; hair 2-X with 4-6
branches; 10 hair tufts in ventral brush, 2-3 tufts proximad of grid; saddle
covered with short rows of fine spicules on lateral surface; anal papillae blunt,
slightly shorter than siphon.
TYPE DATA. Holotype female, Kampong Sireh, Selangor, WESTERN
MALAYSIA, 9 Oct. 1952, J. A. Reid; and paratype female, P. Blakang Mati,
SINGAPORE, 22 April 1955, D. H. Colless, in the British Museum.
DISTRIBUTION. Specimens examined: WESTERN MALAYSIA, Selan-
gov, Rantau Panjang 4 males and 3 females with associated larval and pupal
skins and 1 male. Other Distribution. As in type data.
TAXONOMIC DISCUSSION. A. fvanciscoiis easily recognized by the
presence of broad scales completely covering the mesonotum, absence of pre-
scutellar bristles, presence of broad white scales on the postcoxal membrane,
10 Contrib. Amer. Ent. Inst., vol. 5, no. 4, 1970
and the subapical pale bands on the anterior surface of the femora. The most
marked difference in the male terminalia is the presence of 15-18 moderately
long flattened bristles on the basal mesal lobe.
The larvae resemble iyengari in general appearance but can easily be
separated by the presence of 15-25 comb scales arranged in two rows and head
hair 10-C being single. In iyengari there are 5-10 comb scales in a single row
and head hair 10-C is double.
BIOLOGY. Mattingly (1959: 2, 43) states that Dr. E. E. McClure,
U.S. Army Medical Research Unit, obtained eggs that were laid in a bamboo
pot hung in a mango tree about 30 feet from the ground at Rantau Panjang.
Mattingly believes that this species feeds and breeds almost exclusively at
this height. The holotype female was collected in a trap with human bait.
The allotype, two other males and three females were also reared from eggs
laid in bamboo pots at the afore-mentioned height and location (Mattingly 1961:
59) on various dates in January 1959.
AEDES (DICEROMYIA) IYENGARI EDWARDS
Figs. 2, o?; 9, & terminalia; 10, 11, pupa; 13, larva.
Aedes (Skusea) iyengari Edwards 1923, Bull. ent. Res. 14: 4 (?); Brug 1932,
Bull. ent. Res. 23: 78 (L*, P*).
Aedes Skusea) punctissimus Barraud 1928, Indian J. med. Res. 16: 360 (c*,
2)
Aedes (Dendroskusea) iyengari Edwards 1929, Bull. ent. Res. 20: 341.
Aedes (Diceromyia) iyengari Edwards 1932, Genera Insectorum, Fasc. 194:
173; Barraud 1934, Fauna Brit. India, Diptera 5: 273 (o*, 2, L);
Mattingly 1957, Culic. Mosq. Indomalayan Area 2: 13 (P*); Mattingly
1959, Culic. Mosq. Indomalayan Area 4: 38 (o*, 9*, P¥, L*); Mattingly
1961, Culic. Mosq. Indomalayan Area 5: 58 (c'*).
FEMALE. Head, Antenna dark brown, longer than the proboscis,
torus with a patch of dark scales and short fine hairs mesally, flagellomere 1
with a few dark scales, 1.3 length of flagellomere 2, and with basal 0.5 swol-
len; clypeus dark, bare; palpus dark with a few pale scales at apex, 0. 25-0.3
length of proboscis; proboscis dark with a faint subapical pale spot, about 1. 2
length of fore femur; vertex with alternating dark and pale broad decumbent
scale patches, two patches of pale scales separated by a central dark area,
another pale patch on each side with dark scales below, a few long narrow
erect scales posterior to orbital line and numerous short ones on occiput (the
erect scales on occiput vary from dark to pale). Thorax, Anterior mesono-
tum covered with narrow curved brown scales with a few pale scales inter-
mixed, narrow white scales on anterior margin, a small pale spot of narrow
scales above anterior end and another above posterior end of paratergite;
posterior mesonotum with overlapping long, brown, broad scales along poste-
rior portion and extending over and completely covering scutellum, a patch of
similar scales on supraalar areas; anterior promontory, humeral, 2-4 ante-
rior dorsocentral, supraalar and scutellar bristles well developed, 3-4 poorly
developed prescutellar bristles present, others absent; pleural integument
dark brown; anterior pronotal lobe covered with white scales, narrow curved
Scales above and broad ones below, and several long bristles; upper posterior
pronotum with narrow curved brown scales and a few white ones posteriorly,
middle with a patch of broad white scales, 3-4 posterior bristles; propleuron
with broad white scales and 4-5 bristles; prosternum bare; upper postcoxal
membrane with a few pale scales; spiracular area bare; postspiracular area
Reinert: Aedes (Diceromyia) in Southeast Asia 11
with 4-6 bristles; subspiracular and hypostigial areas with broad white scales;
paratergite with narrow curved white scales along margin; sternopleuron with
upper and lower broad white scale patches and upper and posterior bristles;
prealar area with an upper patch of bristles and broad white scales; mesepi-
meron with an upper patch of broad white scales over anterior, upper and pos-
terior areas, a patch of upper and 2-3 middle bristles. Legs. Coxae with
white scale patches, fore and mid coxae also with dark scales; trochanters
white scaled; anterior surface of femora with apex white, subapical white band
and various other white scale patterns (see figures); fore tibia with apical pale
band, dorsobasal pale spot and 5-6 dorsolateral pale spots over rest of sur-
face, mid tibia with apical pale band and 5 dorsolateral pale spots, hind tibia
with apical lateral pale scales, dorsobasal pale spot and 5-6 dorsolateral pale
spots; fore tarsus with tarsomere 1 having pale scales at apex and base and a
pale band at about middle, tersomeres 2 and 3 with a few dorsobasal pale
scales, mid tarsus with tarsomere 1 having basal and middle pale bands and a
dorsoapical pale spot, tarsomere 2 with some dorsal pale scales at base,
middle and apex, tarsomere 3 with a few dorsobasal pale scales, hind tarsus
with tarsomere 1 having a basal and 2 middle pale bands and a few apical pale
Scales, tarsomere 2 with a few pale dorsal scales at base, middle and apex,
tarsomere 3 with a few dorsobasal scales. Wing. Dorsal veins covered with
broad brown scales with a patch of white scales at the base of the costa; broad
brown scales along posterior margin of wing above fringe scales; alula with
narrow brown scales on fringe; 1-2 remigial bristles. Halter. With pale stem
and knob brown scaled with a few pale scales ventrally. Abdomen. Terga
with brown and pale markings, tergum I with a dorsobasal pale spot and a
dorsoapical pale fringe, terga II-IV each with lateral pale bands curving onto
dorsum and meeting at base in middle, also with 2 admedian pale spots near
apical margin (on Some specimens the bands and admedian pale spots are in-
distinct on terga II-IV), terga V-VII each with lateral pale spots and 2 adme-
dian pale spots near apical margin; posterior margins of terga and sterna
fringed with golden hairs (hairs more numerous on sterna); segment VII com-
pletely retracted into segment VII.
MALE. Similar to female in general habitus. Head. Palpus with
basal bands on terminal 2 segments and a median band on segment 3; palpus
Slightly longer than the proboscis; proboscis with an indistinct pale narrow
band just past the middle. Tevminalia. Tergum IX bilobed with 15-18 approx-
imately equal bristles on each lobe and covered with fine spicules; basimere
short and broad with a patch of moderately long flattened bristles on tergo-
mesal margin near apex, bristles, with only basal portion flattened, in 2-3 ir-
regular rows extending along tergomesal margin from patch to near base, long
bristles extending along dorsolateral margin from base to apex, a patch of
short bristles on dorsal surface below attachment of distimere, ventral sur-
face covered with scattered short bristles and fine spicules, sternomesal mar-
gin with 2-3 long flattened bristles near base, numerous long scales on lateral
and ventral surfaces with a few on dorsobasal area; distimere short, thickened
in the center and tapering to a long blunt point, dark spiniform long and flat-
tened (over 0.5 length of distimere), attached subapically and extending beyond
apex; basal mesal lobe with 5-6 short bristles on a small apical lobe, rest of
surface and fold connecting it to its mate covered with fine hairlike spicules;
phallosome with aedeagus divided into two lateral plates each with 8-11 teeth;
sternum IX with 4 bristles at the center.
PUPA. Cephalothorax. Hairs 1-3, 7-C single; 4, 5, 10, 12-C single,
forked or 2-3 branched. Abdomen. Hair 1-II-III double; 3-III single and as
long as segment III; 1-V single or double; 5-VI single and equal to or longer
than segment VII; 9- VII single and from 0.5 to equal the length of segment VII;
12 Contrib. Amer. Ent. Inst., vol. 5, no. 4, 1970
9-VIII double, occasionally single. Paddle, With hairlike spicules along outer
(distal) 0.75 and inner (distal) 0.5 margins; paddle hair 1-P single.
LARVA. Head, Hair 4-C with 14-21 branches; 6-C double; 7-C with
6-15 branches; 10-C double; 11-C with 10-14 branches; 12-C with 2-4 branches;
antenna short, without spicules and with hair 1-A single and attached at about
0.66 distance from base. Thorax. Hair 9-M stellate with 5-7 branches; 4,
6-T single. Abdomen. Hair 2-I-VII single; hair 1-VII with 2-4 branches;
comb consisting of 5-10 scales in a single row, scales with a fine fringe from
base to the bluntly pointed apex; siphon index approximately 2. 2-2.4; hair 1-S
single and reaching apex of siphon (minus valves); pecten with 6-14 teeth, each
with denticles along ventral and apical margins; hair 1-X single and twice as
long as the saddle; hair 2-X with 4-6 branches; 9-10 hair tufts in ventral
brush, 2-3 tufts proximad of grid; anal papillae blunt and slightly longer than
the siphon.
TYPE DATA, A. iyengari Edwards holotype female, No. 1686,
Meenglas, Jalpaiguri, INDIA, 6 Aug. 1921, M. O. T. Iyengar in British
Museum; paratype female same data, in Iyengar Collection; A. punctissimus
Barraud cotypes male and female; Karwar, INDIA, Sept. 1921, P. J. Barraud,
said to be in the Malaria Institute of India (Barraud 1928: 361).
DISTRIBUTION. Specimens examined: THAILAND, Chiang Mazi, Doi
Sutep 10 whole larvae, Oct. 1955, Thurman. INDIA, Assam, Dibrugarh 3
whole larvae , 8 July 1943, D. E. Hary. INDONESIA, Java 3 males, 6 fe-
males, 1 male and 1 female with associated pupal and larval skins, and 3
pupal and larval skins. INDIA 1 male and 2 females. Other Distribution,
INDIA, Calcutta, Matiabruz, Garden Reach (Barraud 1934: 275); Kidderpore
(Mattingly 1959: 39). INDONESIA, Djakarta (Wijono 1962); Sumba (Bonne-
Wepster 1954: 93); Bandoeng, Purmerend Island (Mattingly 1959: 39). BURMA,
Rangoon (Barraud 1934: 275). THAILAND, Chiang Mai, Doi Pui Mt. (Scanlon
and Esah 1965: 143).
TAXONOMIC DISCUSSION. The adult habitus of zyengari is very sim-
ilar to that of scanloni and is discussed under that species. The male termi-
nalia has similarities to that of fvanciscoi. The major differences from
franciscoi are: basimere with 2-3 long bristles near base of the sternomesal
margin; distimere shorter and apical 0.5 more tapering, spiniform longer and
flatter; and basal mesal lobe with 5-6 short bristles attached to a small apical
lobe. The larva resembles franciscoi but can be easily separated by the pres-
ence of only 5-10 comb scales arranged in a single row while franciscoi has
15-25 comb scales in two rows.
BIOLOGY. Brug (1932: 79) collected larvae in bamboo stumps from
Java at about sea level. Bonne-Wepster (1954: 93) records the larvae breed-
ing in bamboo stumps and tree holes. Scanlon and Esah (1965: 143) report
iyengari biting man at an altitude of 1,000 feet in the forest fringe or in scrub
areas near Doi Pui, Chiang Mai, Thailand.
AEDES (DICEROMYIA) PLATYLEPIDUS KNIGHT AND HULL
Figs. 3, “9; 10, & terminalia.
Aedes (?) platylepidus Knight and Hull 1951, Pacif. Sci. 5: 201 (2); Knight and
Hull 1953, Pacif. Sci. 7: 480 (?L).
Aedes (Diceromyia) platylepidus, Mattingly 1959, Culic. Mosq. Indomalayan
Area 4:43 (9, ?L).
FEMALE. Head. Antenna brown, slightly shorter than the proboscis,
torus with a large patch of broad silvery scales mesally, flagellomere 1 pale,
Reinert: Aedes (Diceromyia) in Southeast Asia 13
with a few dark scales and slightly longer than flagellomere 2; clypeus dark,
bare; palpus dark with apical portion swollen, about 0. 2 length of proboscis;
proboscis dark, approximately equal to fore femur; vertex with broad over-
lapping blackish-brown scales and an anteromedian patch of broad silvery
scales, a patch of pale translucent scales at extreme lower lateral margin,
numerous short black erect scales confined to occiput. Thorax. Anterior and
posterior mesonotum covered with overlapping broad blackish-brown scales
and extending over and completely covering scutellum (mesonotal scales pos-
sess a metallic luster); anterior promontory, humeral, supraalar and scutel-
lar bristles well developed, others absent; pleural integument dark brown;
anterior pronotal lobe covered with bristles and broad brown scales; posterior
pronotum with 3 posterior bristles; propleuron covered with a patch of broad
silvery scales and 1 bristle; prosternum covered with broad silvery scales;
paratergite,spiracular, subspiracular and hypostigial areas bare; postspiracu-
lar area with 2-3 small bristles; a large patch of overlapping broad silvery
scales covering upper and lower sternopleural and anterior, lower, upper and
posterior mesepimeral areas, 1 small golden posterior bristle on sterno-
pleuron and a few golden upper bristles on meSepimeron. Legs, Coxae with
pale silvery scales, mid coxa also with some brown scales; trochanters with
pale and a few dark scales; fore and mid femora dark with a few pale scales at
apex, hind femur with basal 0.6 and apex pale scaled; tibiae dark; fore tarsus
dark, mid and hind tarsomere 1 with a pale dorsobasal spot, hind tarsomere
3 with a pale spot along most of dorsal surface (a few dark scales on dorsal
surface at base and apex). Wing. Dorsal veins mostly covered with broad
brown scales, narrow scales on R., Rg, Rg and median veins; small brown
scales along posterior margin of wing above fringe scales ; alula with narrow
brown scales along fringe; 1 remigial bristle. Halter, With pale stem and
knob brown scaled. Abdomen. Terga with brown and pale markings, terga
II- VI with basolateral pale translucent bands extending a short distance onto
dorsum, tergum VII with a dorsobasal pale translucent band; terga with a few
hairs along posterior margins and sterna fringed with numerous hairs; seg-
ment VOI completely retracted into segment VII.
MALE. Similar to female in general habitus. Head, Antenna about
0.6 length of proboscis; palpus dark, 0.6 length of proboscis; proboscis down-
curved at about middle, with a few pale translucent scales ventrally on apical
0.25, longer than fore femur. Legs. Femora without pale scales at apex;
tarsi dark except for a pale anterobasal spot on hind tarsomere 1. Termina-
lia. Tergum IX with lateral lobes covered with fine hairlike spicules, each
lobe with 2 long scales and 5-6 bristles mesally; basimere with a large patch
of flattened bristles along tergomesal margin, those on basal portion longer,
dorsal surface with a number of short scattered bristles and a number of long
ones along lateral margin and apex, sternomesal margin covered with a long
patch of flattened bristles, ventral and lateral surfaces covered with scales;
distimere with basal 0.5 thickened and apical 0.5 narrow and curved with apex
blunt, spiniform about 0.5 length of apical narrow portion of distimere, dark,
thick, long, curved and attached at about middle of distimere, basal mesal
lobe covered with fine hairlike spicules; phallosome with aedeagus long and
divided into 2 lateral plates each with 8-10 teeth, basal piece large; proctiger
long; sternum IX covered with tiny spicules.
PUPA. Unknown.
LARVA. Not certainly known. Knight and Hull (1953: 480) provision-
ally attributed 3 whole larvae, 2 from Irahuan River (3 miles inland), Palawan
and 1 from Cape Melville, Balabac to this species or Heizmannia scintillans
Ludlow. Mattingly (1957: 32) states the the larvae do not fit Heizmannia very
well.
14 Contrib. Amer. Ent. Inst., vol. 5, no. 4, 1970
The larvae have head hairs 4 single, 7 single or double and siphon
hair 1 inserted before, near or beyond apical pecten tooth which does not com-
pare favorably with the other species of Aedes (Diceromyia) from Southeast
Asia which have head hairs 4 and 7 multiple and stellate and siphon hair 1 al-
ways beyond apical pecten tooth.
The larvae were collected from a large metal packing container in
the jungle and from a large, thorny palm frond lying on the ground at the edge
of a mangrove area which is more like some species of Stegomyia than the
bamboo and tree hole breeding Diceromyia.
TYPE DATA. Holotype female, No. 810.8, Puerto Princesa, Pala-
wan, PHILIPPINES, 24 May, 1945, J. L. Laffoon and Dr. D. R. Johnson;
paratype female (fragment), No. 1620.1, Cape Melville, Balabac, 23 June
1945, J. L. Laffoon; and paratype female (terminalia on slide), No. 1608. 2,
Cape Melville, Balabac; PHILIPPINES, 23 June 1945, D. R. Johnson, in U.S.
National Museum. Paratype female, No. 1608.2, in K. L. Knight collection.
DISTRIBUTION. Specimens examined: PHILIPPINES, Palawan,
holotype female, Balabac, 3 female paratypes, Basilon, Isabella 1 male and 1
female.
TAXONOMIC DISCUSSION. A. platylepidus possesses a number of
characters that differ from most Diceromyia in the Oriental region. The most
marked differences are: the scale pattern of the vertex, absence of scales on
the hypostigial, subspiracular, posterior pronotal, and paratergite areas;
presence of scales on the prosternum, and reduction of sternopleural bristles.
The primary differences in the male terminalia are: tergum IX with
two lateral lobes, each bearing 2 long scales and 5-6 bristles mesally; pres-
ence of flattened bristles along the sternomesal margin on basimere; large
basal piece of phallosome; and a long proctiger. The male palpus also is
shorter than in other species.
Eiven with these differences there exists a number of major charac-
ters that fit the definition of the subgenus and they are: simple tarsal claws
of the female, male antenna with hairs directed mainly dorsally and ventrally,
male palpus with apical 2 segments very short, and structure of the male ter-
minalia, :
Unfortunately, the larval and pupal skins from the adults mentioned
in the distribution could not be found.
BIOLOGY. The holotype was collected as a larva from a small tree
hole along the coast, the paratypes were also collected as larvae, one ina
fallen coconut spathe and the other from a depression in a log in a mangrove
area. One male and 1 female were collected (by K. L. Knight, 27 September,
1945) as larvae from bamboo stakes.
AEDES (DICEROMYIA) PUNCTIPES EDWARDS
Aedes (Skusea) punctipes Edwards 1921, Bull. ent. Res. 12: 77 (9).
Aedes (Diceromyia) punctipes Edwards 1932, Genera Insectorum, Fasc. 194:
174; Barraud 1934, Fauna Brit. India, Diptera 5: 273 (¢).
FEMALE. Head, Antenna missing; palpus dark, slightly longer than
0. 25 length of proboscis which is dark and equal in length to fore femur; ver-
tex with alternating dark and pale broad decumbent scale patches, two patches
of pale scales separated by a central dark area, another pale patch on each
side with dark scales below. Thorax. Anterior mesonotum covered with nar-
row curved bronzy-brown scales, narrow white scales on anterior margin;
Reinert: Aedes (Diceromyia) in Southeast Asia 15
posterior mesonotum with blackish-brown broad scales along posterior and
extending over and completely covering scutellum, a patch of similar scales
on supraalar areas; mesonotal bristles mostly denuded, probably rather long
and dense; anterior pronotal lobe and posterior pronotum with broad white
scales; some other pleural areas with flat white scales. Legs. Femora brown
scaled each with a preapical pale band, white at apex, and pale on ventral
surface, fore and mid femora with white markings basal to preapical band; ti-
biae with apices pale scaled, fore tibia with 3 pale anterior spots, mid and
hind tibiae each with 4 pale anterior spots; tarsomere 1 of fore tarsus with a
basal pale band and a dorsoapical pale spot, tarsomere 2 with a dorsobasal
pale spot; tarsomere 1 of mid tarsus with a basal pale band and dorsal pale
spots at middle and apex, tarsomere 2 with a dorsobasal pale spot; hind tarso-
mere 1 with pale bands at base and middle and a dorsoapical pale spot, tarso-
mere 2 with dorsobasal and dorsoapical pale spots. Wing. Dorsal veins
covered with broad brown scales but a few longer and narrower scales toward
tips of the veins and along lower margin of cubitus,, a patch of pale scales at
base of costa and cubitus. Abdomen. Terga with enw and pale markings,
tergum I with a large pale patch in middle, terga II-VI each with a lateral
pale spot, which is not quite basal and extends a short way onto dorsum; sterna
mostly pale with posterior sterna dark scaled apically; segment VUI complete-
ly retracted into segment VIL.
MALE, PUPA and LARVA. Unknown.
TYPE DATA. Holotype female, Maymyo, Mandalay, BURMA, 11
Dec. 1913, Major Bennett, in the Indian Museum.
DISTRIBUTION. Known only from the type locality.
TAXONOMIC DISCUSSION. No material was available for examina-
tion. The above is based on the original description given by Edwards (1921:
77) and one by Barraud (1934: 273) who had also examined the type specimen.
The description of punctipes is similar to those of zyengari and
scanloni but there are definite differences which are discussed under the taxon-
omic discussion of scanloni.
BIOLOGY. Not known but Edwards (1921: 77) speculates that it is
probably a tree hole breeder.
AEDES (DICEROMYIA) REGINAE EDWARDS
Figs. 4, “9; 8, o terminalia, larva; 10, 12, pupa.
Aedes Skusea) reginae Edwards 1922, Indian J. med. Res. 10: 272 (c).
Aedes (Diceromyia) reginae Edwards 1932, Genera Insectorum, Fasc. 194:
174; Barraud 1934, Fauna Brit. India, Diptera 5: 277 (¢,9,L).
FEMALE. Head, Antenna brown, longer than the proboscis, torus
with a patch of dusky scales and short fine hairs mesally, flagellomere 1 with
a few dusky scales, 1.3 length of flagellomere 2, and with basal 0.5 pale;
clypeus bare; palpus dark scaled, slightly over 0. 25 length of proboscis; pro-
boscis dark scaled, about 1.1 length of fore femur; vertex with alternating
dark and pale broad decumbent scale patches, two patches of pale scales sep-
arated by a central dark area, another dark patch on each side with pale scales
below, a number of long brown erect scales posterior to orbital line (a few
scattered over rest of vertex) and numerous ones on occiput. Thorax. Ante-
rior mesonotum covered with narrow curved golden-brown scales and with
narrow white scales on anterior margin; posterior mesonotum with similar
scales; scales by posterior dorsocentral, prescutellar and supraalar bristles
somewhat longer and broader but still narrow, a small spot of narrow curved
16 Contrib. Amer. Ent. Inst., vol. 5, no. 4, 1970
white scales on ante-alar areas; anterior promontory, humeral, anterior and
posterior fossal, acrostichal, anterior and posterior dorsocentral, supraalar,
prescutellar and scutellar bristles well developed; scutellum with lobes cover-
ed with broad dark brown scales; pleural integument pale yellow; anterior pro-
notal lobe covered with scattered white narrow scales and several bristles;
posterior pronotum with narrow brownish scales, middle with a patch of nar-
row white scales and 4 posterior bristles; propleuron with broad white scales
and 4-5 bristles; prosternum, hypostigial area, postcoxal membrane and spi-
racular area with a few broad white scales; paratergite with a few pale narrow
curved scales along margin; sternopleuron with upper and lower small patches
of broad white scales and upper and posterior bristles; prealar area witha
patch of bristles and a few broad white scales; mesepimeron with a small
patch of broad white scales over anterior and posterior areas, a patch of upper
and 1-3 middle bristles. Legs. Coxae with white scale patches, anterior
coxa also with a few dark scales; trochanters white scaled; femora mainly
brown scaled, hind femur with a pale scaled area on anterior lower 0.5 not
reaching apex; tibiae brown scaled with apical pale scales; tarsi with tarso-
meres dark. Wing. Dorsal veins covered with brown scales, broad scales
on costa, subcosta, Ry and Ry,5 and cubitus, rest of veins covered with nar-
row scales; small scales along posterior margin of wing above fringe scales;
alula with narrow brown scales along fringe; 2-3 remigial bristles. Halter.
With pale stem and knob brown scaled. Abdomen. Terga brown scaled with
pale markings, terga II-VII with basolateral pale bands which are broader on
anterior sterna and may completely cover sterna II and III; posterior margins
of terga and sterna fringed with golden hairs which are more numerous on the
latter; segment VIII completely retracted into segment VIL.
MALE. Similar to female in general habitus. Head, Palpus brown
scaled, slightly longer than the proboscis. Terminalia. Tergum IX bilobed
with 11-12 bristles on each lobe and covered with fine spicules, basimere
short and broad basally, with moderately long bristles covering most of dorsal
surface and long ones along lateral margin to apex, dorsal and ventral surfaces
covered with small spicules, sternomesal margin with a few scattered short
bristles and an apical short flattened lobe with 1 long bristle and several short
hairs, numerous scales on ventral and lateral surfaces; distimere long, nar-
row and curved with a short dark spiniform attached subapically and extending
beyond apex; basomesal lobe covered with fine hairlike spicules and with an
apical patch of 8-10 moderately long bristles; phallosome with aedeagus divid-
ed into two lateral plates each with 8-9 teeth; sternum IX with 7-8 bristles at
the center.
PUPA. Cephalothorax. Hairs 2-4, 7, 10, 11-C single; 1, 5-C single
or double. Abdomen. Hair 1-II-V single; 3-III single, longer than length of
segment III; 5-VI single, equal to the length of segment VII; 9-VII single,
slightly over one-half length of segment VI; 9- VUI with 2-3 branches, slightly
longer than segment VII. Paddle. With small spicules along outer (distal
0.5 and inner (distal) 0.5 margins; paddle hair 1-P single or occasionally
forked.
LARVA. Head, Hair 4-C with 5-15 branches; 6-C single; 7-C with
5-13 branches; 10-C double; 11-C with 5-18 branches; 12-C with 3-4 branches;
antenna short, without spicules and with hair 1-A single and attached at about
0.65 distance from base. Thorax. Hair 9-M stellate with 3-4 branches; hair
4-T single or double and 5-T with 2-3 branches. Abdomen. Abdominal seg-
ments with numerous stellate hairs; hair 2-I-VIII with 1-4 branches, usually
2-3 branched; hair 1-VIII with 2-4 branches; comb consisting of 7-10 scales
in one row, scales with fine fringe along margins and a bluntly pointed apex;
Siphon index approximately 1.4-1.6; hair 1-S with 2-3 branches and not
Reinert: Aedes (Diceromyia) in Southeast Asia 17
reaching apex of siphon; pecten with 4-10 teeth with denticles along ventral and
apical margins; hair 1-X with 3-5 branches and about 1. 2 length of saddle;
hair 2-X with 4-7 branches; 10-11 hair tufts in ventral brush, 1-2 tufts proxi-
mad of grid; saddle covered with short rows of fine spicules on lateral sur-
face; anal papillae broad and blunt, approximately 0.75 length of siphon.
TYPE DATA, Types 1 male and 3 females, Colombo, CEYLON,
1913, James, in the British Museum.
DISTRIBUTION. Specimens examined: CEYLON, Colombo 3 females,
2 males with associated larval and pupal skins. INDIA, Nicobar Islands 1 fe-
male. INDIA, mainland 1 female. Other Distribution. INDIA, Kharghpur,
Bengal-Nagpur Rly. (Barraud 1934: 277).
TAXONOMIC DISCUSSION. A. rveginae is very similar to micropterus
in the adult habitus and the male terminalia, It has the pleural integument
pale yellow, the apices of all femora dark scaled and the lateral surface of the
head pale scaled compared to micropterus in which the pleural integument is
dark brown, the apices of all femora with lateral white spots and the lateral
surface of the head dark scaled. The male terminalia of reginae also differ
from micropterus in having the baSimere shorter, wider and with more nu-
merous bristles along the lateral margin. The larvae are very similar to
micropterus but differ, according to Mattingly (1959: 38), in having the inner
mouthbrush setae pectinate, papilliform appendage of antenna short and undif-
ferentiated, and hair 2-X with 4-7 branches. A. micropterus has the mouth-
brushes without pectinate setae, papilliform appendage of antenna long, narrow
and partly differentiated into two, and hair 2-X with 3-4 branches. A. reginae
differs from the other Oriental species in having numerous stellate hairs on
the abdominal segments.
BIOLOGY. The larvae of the type specimens were collected from
tree holes (Edwards 1922: 272).
AEDES (DICEROMYIA) SCANLONI n. Sp.
Figs. 5, “9; 9, o& terminalia.
FEMALE. Head, Antenna dark brown, longer than the proboscis,
torus with a patch of dark scales and short fine hairs mesally, flagellomere 1
with a few small dark scales and 1.3 length of flagellomere 2; clypeus dark,
bare; palpus dark with apical pale band which is wider mesally, slightly longer
than 0. 25 length of proboscis; proboscis dark brown, slightly longer than fore
femur; vertex with alternating dark and pale broad decumbent scale patches,
two patches of pale scales separated by a central dark area, another pale
patch laterally on each side with dark scales below, a few dark long narrow
erect scales posterior to orbital line and numerous short narrow ones on oc-
ciput which also has a few pale erect scales. Thorax. Anterior mesonotum
covered with narrow curved brown scales, narrow curved white scales around
anterior margin, a small white patch above paratergite and one on the anterior
margin of supraalar area; posterior mesonotum with overlapping long brown
broad scales along posterior and extending over and completely covering scu-
tellum, a patch of similar scales on supraalar area; anterior promontory,
humeral, 2-4 anterior dorsocentral, supraalar and scutellar bristles well de-
veloped, 3-4 poorly developed prescutellar bristles present, others absent;
pleural integument dark brown; anterior pronotal lobe covered with white scales
and several long bristles, narrow curved scales above and broad ones below;
upper posterior pronotum with narrow curved brown scales and a few white
ones posteriorly, middle with a patch of broad white scales, 4 posterior bris-
tles; propleuron with broad white scales and 3-4 bristles; prosternum bare;
18 Contrib. Amer. Ent. Inst., vol. 5, no. 4, 1970
postcoxal membrane and spiracular area bare; postspiracular area with 4
bristles; subspiracular and hypostigial areas with broad white scales; para-
tergite with narrow curved white scales; sternopleuron with upper and lower
broad white scale patches and upper and posterior bristles; prealar area with
a patch of bristles and a few broad white scales; mesepimeron with a patch of
broad white scales over anterior and posterior areas, a patch of upper and 2
middle bristles. Legs. Coxae with white scale patches, fore and mid coxae
also with dark scales; trochanters white scaled; anterior of femora with a few
pale scales laterally at apex, a subapical pale band and various other pale
scale patterns (See figures); tibiae each with 4 dorsolateral evenly spaced pale
spots and apical pale scales; fore tarsus with tarsomere 1 having pale dorsal
scale patches at base, apex and middle, tarsomeres 2-4 with a few dorsobasal
pale scales; mid tarsus with tarsomere 1 having pale spots at base, apex and
2in middle, tarsomeres 2 and 3 with a few dorsobasal pale scales; hind tarsus
with tarsomere 1 having pale bands at base and 2 in middle, and a few pale
apical scales, tarsomeres 2-4 each with a dorsobasal pale spot and a few dor-
soapical pale scales. Wing. Dorsal veins covered with broad brown scales
with white ones at bases of costa, radius and cubitus (in Some specimens 1 or
2 pale scales at base of anal vein also); small brown scales along posterior
margin of wing above fringe scales; alula with narrow to broad brown scales
along fringe and a short second row above; 1-2 remigial bristles. Halter,
With pale stem and knob brown scaled with a few pale scales ventrally. Ab-
domen. Terga with brown and pale markings, tergum I with a large dorsal
pale patch from base to apex, terga II-VI each with a lateral pale patch, ter-
gum II with a large dorsobasal pale patch, terga V-VI with 2 admedian pale
spots near apical margin in the holotype (Some paratypes also with 2 admedian
pale indistinct spots on terga III, IV and VIL); posterior margins of terga and
sterna fringed with golden hairs which are more numerous on the latter; seg-
ment VIII completely retracted into segment VIL.
MALE. Similar to female in general habitus. Head, Palpus with
broad basal pale band on terminal segment, segment 4 with a few dorsobasal
pale scales and a median pale band on segment 3; palpus slightly longer than
the proboscis; proboscis with an indistinct pale narrow band just past middle.
Terminalia. Tergum IX bilobed with 8-10 more or less equal bristles on each
lobe and covered with fine spicules; basimere short and broad with a patch of
moderately long flattened bristles along upper 0.5 of tergomesal margin,
shorter flattened bristles extending along tergomesal margin to base in 2-3 ir-
regular rows, long bristles extending along dorsolateral margin from base to
apex, ventral surface covered with scattered short bristles and fine spicules,
sternomesal margin with 6-9 moderately long flattened bristles near base,
numerous long scales on lateral and ventral surfaces with a few on dorsobasal
area; distimere short, thickened in the center and tapering to a blunt point,
spiniform dark, long and flattened (over 0.5 length of distimere), attached
sub- apically and extending beyond apex; basal mesal lobe covered with fine
hairlike spicules and with numerous short flattened bristles on apical portion;
phallosome with aedeagus divided into two lateral plates each with 10-11 teeth;
sternum IX with normally only 2 bristles at the center (paratype NB 681 with
4 bristles). :
PUPA and LARVA. Unknown.
TYPE DATA. Holotype female, U-50B, Nakhon Ratchasima, THAI-
LAND, 29 May 1963, Resting Collection; allotype male, Don Mung Road,
Krung Thep, THAILAND, 30 Aug. 1955, M.N.R.; 1 female paratype, U-50
with same data as holotype; 1 male terminalia, paratype, on slide and 2 fe-
male paratypes with same data as allotype; 1 male paratype, NB1135, Pak
Kret, Nonthaburi, THAILAND, 23 Aug. 1964, resting in house; 1 male
Reinert: Aedes (Diceromyia) in Southeast Asia 19
paratype, NB681, from an island in the Chao Phraya River, Pak Kret, Nontha-
buri, THAILAND, 16 June 1964, light trap collection; 1 male terminalia
(adult lost) paratype, T-5033, slide 2317, Udornthani, Udon Thani, THAI-
LAND, 15 Sept. 1962, light trap. All type specimens deposited in the U.S.
National Museum.
DISTRIBUTION. In THAILAND from the following changwats: Krung
Thep, Nakhon Ratchasima, Nonthaburi, and Udon Thani.
TAXONOMIC DISCUSSION. A. scanloni is very similar to iyengari
and punctipes in the adult habitus. The primary differences from iyengari
are: presence of pale scales on the wing at the bases of the radius and cubitus;
postcoxal membrane bare; and the tergal markings, especially tergaI and II
which have large dorsal pale areas. The male terminalia also has a number
of marked differences from iyengari. These differences are: tergum IX with
8-10 bristles; basimere with longer patch of flattened bristles on tergomesal
margin, without a patch of short bristles on dorsal surface below attachment
of distimere, with basal mesal lobe covered with approximately 25-35 short
flattened bristles, sternomesal margin with 6-9 moderately long bristles near
base; and with normally only 2 bristles on sternum IX,
The primary differences from punctipes are: presence of apical pale
band on palpus; legs with more extensive pale markings; pale scales on the
wing at the base of the radius, and terga V-VI with 2 admedian pale spots near
apical margin.
This species is named for LTC John E. Scanlon in recognition of his
valuable work on mosquitoes in Southeast Asia.
BIOLOGY. Adults were collected in light traps and resting in houses.
AEDES (DICEROMYIA) WHARTONI MATTINGLY
Figs. 6, &9; 7, o terminalia; 10, 12, pupa; 15, larva.
Aedes (Diceromyia) whartoni Mattingly 1965, Culic. Mosq. Indomalayan Area
6: GhAe*,; P* Li).
FEMALE. Head, Antenna dark brown, longer than the proboscis,
torus with some short fine hairs mesally, flagellomere 1 with a few small dark
scales on basal 0.5 and 1.3 length of flagellomere 2; clypeus dark, bare; pal-
pus dark, slightly over 0. 25 length of proboscis which is dark and slightly lon-
ger than fore femur; vertex with alternating dark and pale broad decumbent scale
patches of variable size, 2 small pale patches separated by a central dark area,
another pale patch on each side with dark scales below, a few erect scales post-
erior to orbital line and numerous ones on occiput. Thovax, Anterior mesono-
tum, covered with long narrow curved blackish-brown scales with narrow white
scales around anterior margin, a patch of broad white scales above and extending
down over paratergite; posterior mesonotum with overlapping long black broad
scales along posterior and extending over and completely covering scutellum,
Similar scales extending forward and covering supraalar area; anterior promon-
tory, humeral, supraalar and scutellar bristles well developed, others absent;
pleural integument dark brown; anterior pronotal lobe with some bristles and
covered with broad white scales; upper and middle posterior pronotum covered
with a patch of broad blackish-brown scales, 4-5 posterior bristles; propleuron
covered with broad white scales and 2-3 bristles; prosternum bare; postcoxal
membrane and spiracular area bare; postspiracular area with 4-5 bristles; Sub-
Spiracular and hypostigial areas with broad white scales; paratergite com-
pletely covered with overlapping broad white scales; sternopleuron with upper
and. lower broad white scale patches and upper and posterior bristles; prealar
20 Contrib. Amer. Ent. Inst., vol. 5, no. 4, 1970
area with a patch of bristles; mesepimeron with a patch of broad white scales.
over anterior and posterior areas; a patch of upper bristles present, other
bristles absent. Legs. Coxae with white scale patches, fore coxa also with
brown scales; trochanters white scaled; femora each with a subapical white
band, a very narrow basal pale band and white scales at apex; tibiae brown
and with a few dorsoapical pale scales; fore and mid tarsomere 1 each with a
dorsobasal pale spot, mid tarsomeres 2-4 with a few dorsobasal pale scales,
hind tarsomeres 1 and 2 each with a basal pale band, hind tarsomeres 3, 4
and usually 5 with a dorsobasal pale spot. Wing. Dorsal veins covered with
broad brown scales and a patch of white scales at base of the costa; small
broad scales along posterior margin of wing above fringe scales; alula with
broad brown scales along fringe and a second row above; 1 remigial bristle.
Halter. With pale stem and knob brown scaled. Abdomen. Terga with dark
brown and white markings, terga I-VI dark dorsally, terga II-VI each with a
lateral white spot near basal margin; posterior margins of terga and sterna
fringed with golden hairs (hairs more numerous on sterna), basally sterna
II-VI with prominent white bands; segment VUI completely retracted into seg-
ment VU.
MALE. Similar to female in general habitus. Head. Palpus dark,
slightly longer than the proboscis. Abdomen, Terga VI-VII with lateral
white spots extending onto dorsum. Terminalia. Tergum IX bilobed with 9-15
bristles on each lobe, and covered with fine spicules; basimere short and
broad basally, with numerous long bristles on dorsal surface and along the
tergomesal margin, a number of longer bristles on upper lateral surface and
at apex, a few short bristles along sternomesal margin and ventral surface,
also a patch of 10-12 long bristles near base of sternomesal margin, a few
short hairs at apex mesally, numerous dark scales on ventral, lateral and
laterobasal portion of dorsal surface, ventral and dorsobasal surfaces covered
with small spicules; distimere long and curved with a lateral flaplike structure
just past center in most Specimens, spiniform dark, short, attached subapical-
ly and extending beyond apex; basal mesal lobe with 5-9 moderately long bris-
tles on a small apical lobe, rest of surface and narrow fold connecting it to
its mate covered with fine hairlike spicules; phallosome with aedeagus divided
into two lateral plates each with 9-11 teeth; sternum IX with 1-2 stout bristles
at the center.
PUPA. Cephalothorax, Hairs 1, 2-C double; 3-C with 2-3 branches;
4, 5, 7, 12-C single or double; 10-C single or 3-branched; trumpet moderate-
ly long. Abdomen. Hair 1-II-VI single or double; 1-VII single; 3-III single,
longer than the length of segment III; 5-VII single, slightly less than 0.5 the
length of segment VI; 9-VII single or with 2-4 branches, longer than the length
of segment VII. Paddle. With tiny spicules along proximal 0.5 of outer mar-
gin; paddle hair 1-P with 2-5 branches.
LARVA. Head, Hair 4-C large, stellate with 11-21 branches; 6-C
double; 7-C with 3-9 branches; 10-C single or with 2-3 branches; 11-C stel-
late with 7-12 branches; 12-C with 3-4 branches; antenna long, with a few
Spicules on basal 0.5, hair 1-A double and attached at about 0.65 distance
from base. Thovax. Hair 9-M stellate with 3-4 branches; hairs 4, 5-T single.
Abdomen. Hair 2-I-VIII single, hair 1-VII single, seldom double; comb con-
sisting of 10-14 scales in 2 irregular rows, scales with sharp pointed apex
and fringe on basal 0.75 of lateral margins; siphon index approximately 3. 6-
4.1; siphon covered with minute spicules and apical 0.5 conical; hair 1-S sin-
gle, not reaching apex of siphon; pecten with 9-14 teeth, with denticles along
the ventral and apical margins; hair 1-X single, slightly longer than saddle;
hair 2-X with 2-3 branches; 8-9 hair tufts in ventral brush, 1-2 tufts proxi-
mad of grid; saddle covered with short rows of fine spicules on lateral surface;
Reinert: Aedes (Diceromyia) in Southeast Asia 21
anal papillae blunt, short and approximately 0.3-0.5 length of siphon.
TYPE DATA. Holotype male with associated larval and pupal skins,
No. 0923/9, Ulu Gombak, WEST MALAYSIA, 20 Nov. 1958, W. W. Macdonald,
in British Museum.
DISTRIBUTION. Specimens examined from the following changwats
in THAILAND: Lampang 2 females and 1 male with associated larval and pupal
skins, 1 female with associated pupal skin, 1 whole larva; Nan 1 female with
associated larval and pupal skins; Phangnga 2 females and 3 males with asso-
ciated larval and pupal skins, 8 females and 5 males with associated pupal
skins, 1 female and 2 males and 12 whole larvae; Ranong 2 females and 3
males with associated larval and pupal skins, 2 males with associated pupal
skins and 1 male; Tak 1 female and 1 male. WESTERN MALAYSIA, Selangor,
The Gap 1 female with associated pupal skin, 1 male and 4 females, Ulu Gom-
bak 1 female with associated larval and pupal skins, 1 male and 2 females.
TAXONOMIC DISCUSSION. A. whartoni is a dark species and easily
separated from the other Oriental species of Diceromyia by the absence of
anterior dorsocentral and prescutellar bristles, presence of a patch of broad
white scales above and extending over the paratergite, broad blackish- brown
scales on the posterior pronotum, and lateral tergal markings of the abdomen.
The male terminalia resembles those of reginae, micropterus and periskelatus
but can be readily separated by the presence of a patch of 10-12 long bristles
on the sternomesal margin of the basimere near the base and the flaplike struc-
ture (when present) on the distimere. The larvae are easily recognized from
the other species of Oriental Diceromyia by the presence of the long antenna
and siphon, It is like franciscoi in having two rows of comb scales but differs
in having only 10-14 scales compared to 15-25 scales in fyanciscot,
BIOLOGY. Immatures in Thailand have been collected from fresh,
colored water in large and small split bamboo, -bamboo internodes, and bam-
boo stumps; 1-2 meters above the ground; in mountain, hilly and valley ter-
rain; in partial and heavy shade; in secondary rain forests and secondary bam-
boo groves; and at an altitude of 100 to 1,600 meters (most commonly collect-
ed at altitudes of 200-300 meters in secondary rain forests). One collection
of larvae was taken from a hole in a log lying on the ground. Mattingly (1965:
66) records A, whartoni breeding in fallen bamboos in Malaya.
In Thailand immatures were collected in association with the follow-
ing species of mosquitoes: Aédes albolineatus, albopictus and niveus group;
Anopheles asiaticus; Armigeres durhami; Culex species; Heizmannia species;
Orthopodomyia albipes and andamanensis; Toxorhynchites gravelyi. and
leicestert; Tripteroides avanoides and similis; Udaya argyrurus; and Urano-
taenia bimaculata and lutescens.
ACKNOWLEDGEMENTS
I am grateful to Dr. Botha de Meillon, Responsible Investigator,
Southeast Asia Mosquito Project (SEAMP) and to Dr. Alan Stone, Agriculture
Research Service, U.S. Department of Agriculture, for their many helpful
suggestions and criticisms during the course of the work and preparation of
the final manuscript. Sincere thanks are given to LTC John E. Scanlon, U.S.
Army, E.L. Peyton and Dr. Yiau-Min Huang, all of the Southeast Asia Mos-
quito Project, for their helpful suggestions and encouragement. Dr. P. F.
Mattingly kindly read the manuscript and made many helpful suggestions. The
loan of important material, obtained for me by SEAMP, from Dr. P.F.
Mattingly, Department of Entomology, British Museum (Natural History); Dr.
E.C.C. van Someren, Division of Insect- Borne Diseases, Nairobi, Kenya;
22 Contrib. Amer. Ent. Inst., vol. 5, no. 4, 1970
Dr. T. Ramachandra Rao, Virus Research Centre, Poona, India; The Director,
South African Institute for Medical Research, Johannesburg; Dr. L. Rozeboom,
Johns Hopkins University; and Dr. J. Bonne-Wepster, Instituut voor Tropische
Hygiene en Geographische Pathologie, Amsterdam, The Netherlands, is ac-
knowledged with sincere appreciation. I am particularly indebted to the per-
sonnel of the SEATO Medical Research Laboratory, Thailand, for the collec-
tion and preparation of many valuable specimens. Thanks are extended to Mr.
C. John Lane for his valuable work as curator, Mrs. Elaine R. Hodges for
preparing the immature and male terminalia illustrations, the artists of the
U.S. Army 406th Medical Laboratory, Tokyo, Japan, for preparing the adult
habitus illustrations, and Miss Helle Starcke for typing the manuscript for off-
set reproduction. To my wife, Mollie, I am especially appreciative for her
assistance in typing the drafts.
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APPENDIX: TABLE 1
Species of Aedes (Diceromyia) occurring in the Ethiopian Zoogeographical
Region.
SPECIES FEMALE MALE PUPA LARVA EGG
adersi X X* . K* =
bananea ».4 = © 3 :
Jascipalpis xX* x* x ». Gs -
flavicollis x* x* ».4 ».4 =
furcifer Xx* X* x* X -
ZvYAasset Xx X* = = ‘
taylori x x* D4 x =
zethus xX X* xX X* =
‘e ndicates stage has been described in the literature.
=]
= Indicates a portion of the stage has been figured in the literature.
= Indicates no description or figure.
Reinert: Aedes (Diceromyia) in Southeast Asia 27
APPENDIX: TABLE 2
Species of Aedes (Diceromyia) occurring in the Oriental Zoogeographical
Region.
SPECIES FEMALE MALE PUPA LARVA EGG
franciscot xX** x** K** X** “
LVEngart xX** xX** X** XK i
Ranavensis D4 - ~ - -
micvopterus x xX* g: xX* <
periskelatus x xX* X* X* ss
platylepidus X** X** * X? .
punctipes x - ~ - -
vamachandrai Xx* x* ~ - ‘
reginae X** xX** X** Xe -
scanloni X** X** - - 2
whartoni x** X** Xx Xx e
X = Indicates stage has been described in the literature.
* = Indicates a portion of the stage has been figured in the literature.
** = Indicates a portion of the species is figured in the present paper.
? = Indicates not a positive association.
- = Indicates no description or figure.
an
t
1.0 mm———_——_4
/
/SCO
franc
/yengari
platylepidus
s
ee,
c
a
=
AA}
1S)
w
’
/
wharton
Woy f
ARNON TUelnes
Ky ate
\
Vay hiy
(Ubrl tY
Pay yay
pale ees eos
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wy
NN
MALAYA
francisco/
variable
distimere
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‘ Ua Zn ea) pinefe J
on a 2oeee pet
Wt aces yay
te rgum | X
yi Atte: tht tet
Maen '
fave ryt
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sternum |X
\\ basolateral
\ sclerotization
of proctiger
O.lmm
MALAYA, THAILAND
wharton/
0.05mm
\
10
inner
mouthbrush
seta
1!
CUA EROS)
\
M4
paraproct
basolateral
sclerotization
of proctiger
CEYLON :
reginae
f Ys,
Pv VO Naty ae
RSH , . he) Se
ea tag rt
PANE ese
Nei
sternum |X
basolateral sclerotization
of proctiger
mee Vid yl i
decyl 3
che go,
ME. y JAVA
lyengari
v “yyy
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SOREL
WA
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|
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Vw Wah iY Wy \ ‘i A Sey
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beet)
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THAILAND
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MeN Wyeaava,
A \ MW. abs ] aN
ete: ge Ma, Pr gee My
TAN set yay fea Bh
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CU 2 a RE x, \'
MY WV SS ESIC
=e = Wie
YP ATTAT A ATINERS
fy
paraproct
basolateral sclerotization
of proctiger
paramere
aedeagus
PHILIPPINES
platylepidus
IVY
‘NS
SNR
~
cern {
Min ” i if
hy, f
ibe peal
francisco/
whartoni/
/yengar/
pupal paddles
<
>
<
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<
=
o—
.@)
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D
~
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N
Reinert: Aedes (Diceromyia) in Southeast Asia 43
INDEX
Valid names are set in roman type, synonyms are italicized. The itali-
cized pages are those which begin the primary treatment of the taxon.
Num-
bers in parenthesis refer to the figures illustrating the species in question.
adersi
Aedes
Aedes albolineatus
Aedes albopictus
Aedes niveus
Aedes (Skusea)
Aedimorphus
africana
Anopheles asiaticus
Armigeres durhami
bananea
Culex
Dendroskusea
Diceromyia
fascipalpis
Finlaya
flavicollis
franciscoi
fur cifer
grassel
Heizmannia
Heizmannia scintillans
lyengari
kanarensis
Mansonia
micropterus
Ochlerotatus
Orthopodomyia albipes
Orthopodomyia andamanensis
periskelatus
platylepidus
punctipes
punctissimus
ramachandrai
reginae
scanloni
Stegomyia
taylori
Toxorhynchites gravelyi
Toxorhynchites leicesteri
Tripteroides aranoides
Tripteroides similis
Udaya argyrurus
Uranotaenia bimaculata
Uranotaenia lutescens
whartoni
zethus
“
)
, 8. 19, B1, 27.44. 7, 16, 14, 14),
)
ww
8 OF O1 GO OO >
& 9 e
Ww wr
e ~
bo’
—_
6.1, 10, 22, 35; 19, Bt, (2, 9, £6,
©» frascd. w
no oo
©
[St ~]
. ( ae
DO
=
~ Se Teal, at, at.
1.
1
2) 8, 7, io 27.
2.9 4 6 6 is 1a Be, tay iy.
4, 14, 15, 16, 27. |
0, 12.
, 2. 8. Be te ae
6. 9, 15 Bi, 20, tA. Gy 10, 12):
8.4. 7, 12 15, 27,10: 27, G, oy
ers
, Bs 26.
iL
21
21
21.
at,
24.
21
> 3, , 6, %, 19, 21; 2716. FW; 19, 15)
6.
POR ole Menthe HOC snl ee
:
ae 3
ae ee
ie
orn
Ss
page
mt
No.
No.
(Continued from inside front cover)
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Contributions
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American Entomological Institute
Volume 5, Number 5, 1970
THE TANYMECINI OF THE WEST INDIES
(COLEOPTERA: CURCULIONIDAE)
by
AN HSO/ i Ay
MAY 181970 |
Anne T. Howden
CIBRARIE2
yt
CONTRIBUTIONS
of the
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(Continued on back cover)
THE TANYMECINI OF THE WEST INDIES
(COLEOPTERA: CURCULIONIDAE)"
By
Anne T. Hewden®
ABSTRACT
The Tanymecini of the West Indies are discussed, with the exception of Polydacrys
Schénherr and Pachnaeus Schénherr. The new genus Scalaventer is widespread and in-
cludes the following species: cyrillae, new species, from Jamaica; coccolobae, new
species, from Jamaica; litoreus, new species, from Jamaica; montanus, new species,
from Jamaica; remotus, new species, from Jamaica; subtropicus (Fall), new combina-
tion, from Cuba; caymani, new species, from Grand Cayman; jamaicensis, new species,
from Jamaica; convexifrons, new species, from the Bahama Islands; gelinasus, new
species, from Dominican Republic; valkyrius, new species, from Jamaica; and cubensis,
new species, from Cuba. The new genus Paululusus is endemic to Hispaniola and con-
tains three new species, calypso, hispaniolae, and constanzae. The new genus Para-
dacrys is endemic to the Bahama Islands and contains two new species, spatulatum and
ensiformis. Pandeleteius Schonherr is represented by testaceipes Hustache, endemic to
the Lesser Antilles, and nodifer Champion established on Jamaica. The genus Isodrusus
Sharp is represented by insulanus Howden from the Bahama Islands and guajavus, new
species, from Jamaica. Lectotypes are designated for Pandeleteius subtropicus Fall
and Pandeleteius nodifer Champion.
Biological information is given for many of the species from Jamaica. All teratologi-
cal evidence is recorded, and the zoogeography of the tribe is briefly discussed.
CONTENTS
PM VOGUCTION. pete gee wer et i Pie arta Ge fas ae pa ae
BECUAOI S55 eae ei ee ee a) bao ee ON ah a Bias 2
ey OPE IIAICS Alig’ ylG@eme ee 3
Paunal Gist by Islands 6 os ee ee ee as ears «- $2
Zoeceveorraphic Retatignshins.. 0. foe. be oe Ne ea ear Do
Acknowledgments. ........ nn) Se eae ni etre ee a D0
perature Cue sy bee os ee i a a ae 06
AGC Sod ie: in as gh as eo ates os OA Gate me Gal, Nolan ull a til i Bt
gee. 5 eS es pe ge eas ee ae es tle a as Boe eo ne
This is the sixth paper in a series on Tanymecini.
“Research Associate, Biology Department, Carleton University, Ottawa, Ontario,
Canada.
2 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
INTRODUCTION
The presence of ocular vibrissae is the traditional character for distin-
guishing Tanymecini from other tribes with a mandibular scar and without ocu-
lar lobes. The three new genera herein described, Scalaventer, Paululusus,
and Paradacrys, lack any trace of ocular vibrissae. The separated fore coxae
alone remains to eatablish these genera as tanymecines and to distinguish them
from other Brachyderinae with similar habitus.
It is note worthy that in the continental genera most closely related to these
West Indian genera the condition of the vibrissae varies interspecifically. Of
fifteen species of Isodacrys (some undescribed) the vibrissae are completely
absent in seven species, variable or vestigial in five species, and well devel-
oped in three species. Of the three species of Isodrusus, the vibrissae are
absent in one, vestigial in one, and well developed in one. Of five species of
Pandeleteinus, the vibrissae are vestigial in one and well developed in four,
It is also interesting to note the condition of the wings while comparing
characters of higher taxa. The apterous condition is uniform and stable for
Isodacrys, but does occur sporadically within at least two other well-known
tanymecine genera where it has only specific rank or less. In the very large
genus Pandeleteius at least dentipes Pierce and simplarius Fall have fused
elytra and only rudimentary wings. In the genus Tanymecus specimens of con-
fertus Gyllenhal from one locality may have rudimentary or fully-developed
wings (Champion 1911, pp. 178, 179). In the new genus Paululusus, two spe-
cies have well-developed wings, and one is apterous.
All species discussed in this paper (if not all Tanymecini) have the first
two ventrites of the abdomen fused and the elytra with an interlocking mechan-
ism consisting of a narrow flange on the right elytron which fits into an inter-
nal pocket on the left elytron.
METHODS
Color and color pattern are best observed at lowest power or without a
microscope. Sculpture of the scales is described as it appears at 54x or more.
Setae should be observed in profile as well as dorsally. Length of specimens
is measured from the anterior edge of eyes to apex of the elytra in dorsal view.
Width of the head between the eyes and the width of the beak must be carefully
measured, preferably at high power. ‘‘Interantennal line’’ refers to a trans-
verse ‘‘line’’ between or just caudad of insertion of the antennae on the dorsum
of the beak; like the ‘‘median line’’ it may or may not be visibly modified.
The prothorax is measured for length dorsally along the median line, for
width dorsally at the widest point, and for thickness laterally at the base.
Measurement for relative length of pronotum and prosternum is made from
lateral view.
Males may be externally distinguished from females by the morphology of
ventrite 5. In males the apex of this ventrite is truncate or emarginate, de-
flected and never margined; in females the apex of ventrite 5 is rounded, never
deflected and usually finely margined. When both sexes of a species are at
hand for comparison, the males can be distinguished by their more narrowly
separated fore coxae, narrower head and beak, narrower and straighter elytra,
more convex eyes, and thicker fore femora.
Types in the Howden collection are stored in the Canadian National Collec-
tion.
Howden: West Indian Tanymecini 3
SYSTEMATICS AND BIOLOGY
Key to West Indian Genera of Tanymecini
RR ed so) ME eo: BRM RR YO nae aS A a ma Pen MMC MC CaM ee SR MBER IME Ne 2
Corbels enclosed or semi-enclosed. ........2.6+s-2ece-s-scevee 6
2. Tarsal claws connate. Bahama Islands, Jamaica. ..........
AS ES PP MR IE ig Farin, Mee Ge gk PE armed Isodrusus Sharp (p. 50)
TALSAL Cine EP ae vie cis aa eee aa ck, ee dente eee oa ee ts
3. Ocular vibrissae well-developed. Anterior portion of ventrites 3, 4 and 5
flat and unmodified. Lesser Antilles and Jamaica .........
Sikh. oe ae dBase a ae a ee Pandeleteius Schoénherr (p. 45)
Ocular vibrissae absent. Anterior portion of ventrites 3, 4 and 5 glabrous
BNC CONCAVE WOGtALIY Ge JOR 6ts Sek cc ek. eae A ee ee ee Rk me 4
4. Anterior portion of ventrites 3, 4 and 5 with strong modifications extend-
ing entire width of abdomen. Bahama Islands, Hispaniola, Jamaica,
Cuoan, southern Fioriga, tie. Catornig i secs ee ee es
is eas Sab 1% Aa ee OE ne: a aes ee Scalaventer, new genus (p. 3)
Anterior portion of ventrites 3, 4 and 5 modified with at most a narrow
glabrous area (often medial only) and concavity (medial or entire width)
wWhich.ts always Penlie. Mever CAringtes ore ssc% sa T eae a hae we 5)
5. Fore coxae in both sexes separated by no less than four-fifths the width and
usually more than the width of the antennal club. Eye well separated
from dorsal surface. Aedeagus much thicker than antennal scape. Ba-
PA le ee ae a ge es Paradacrys, new genus (p. 40)
Fore coxae in both sexes separated at most by one-half the width of the
antennal club. Eye very close to dorsal surface. Aedeagus scarcely
thicker, if at all, than antennal scape. Hispaniola .........
ge CEN er TO Se Fo pe RG NTE aE ge Paululusus, new genus (p. 32)
6. Corbels semi-enclosed. Apical emargination of beak keeled. Mandibular
cusp situated at the apex of an anterior projection of the mandible.
Greater Antilles, Guadeloupe. ......... Polydacrys Schoénherr
Corbels enclosed. Apical emargination of beak not marked by a keel or
carina. Mandible not produced. Greater Antilles. ........
iit, Vek a atest ll a gd at MCL hl AR cee ce Ae a ag a Pachnaeus Schoénherr
Scalaventer, new genus
Size small to moderate, usually between 2.3 and 4.5 mm. in length. Body
and legs covered with scales and small setae. Beak of various forms; inter-
antennal line usually unmarked. Apex of beak emarginate, the carina marking
the emargination obsolete to fine. Mandibles extending well beyond apex of
beak. Ventral edge of scrobe usually visible from above. Scrobe of various
Shapes, usually reaching ventral surface or nearly so. Antennal scape without
scales; funicle seven-segmented, the first segment nearly twice as long as the
remaining segments which are equal to each other or nearly so; club moderate
to large, often slightly pedunculate and not completely compact. Head in lateral
4 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
profile transversely swollen between or behind eyes, or not. Eye moderately
small to large in size, large faceted, usually strongly convex and protruding.
Pronotum produced anteriorly over head; anterior margin of prothorax unmodi-
fied, i.e., without ocular vibrissae, teeth or lobes. Elytra with humeral an-
gles well developed. Stria 10 obsolete medially. Fore legs scarcely to con-
spicuously larger than other legs; fore femur swollen or not, without teeth or
serrations on inner edge; fore tibia dentate on inner edge. Fore coxae usually
separated by less than the width of the antennal club. Hind tibia with corbel
open. Tarsal claws free. Abdomen with anterior portion of ventrites 3, 4 and
9 strongly modified across the entire width, i.e., glabrous, deeply, and abrupt-
ly concave, the concavity often delimited posteriorly by a strong carina. Aedea-
gus Slender.
Type-Species. -Scalaventer cyrillae new species.
Scalaventer is masculine and refers to the transverse depressions and ridges
of the ventrites which in many species are not unlike the rungs of a ladder.
These gross modifications of ventrites 3, 4 and 5 are the principal diagnostic
character for the genus.
Discussion. - Of the continental New World genera, Scalaventer is closest
to Pandeleteinus and Isodrusus; of the West Indian genera, Scalaventer is close
to both Paradacrys and Paululusus as well as Isodrusus. In comparison with
all these genera, except Isodrusus, the critical difference is the modification
of the anterior portion of ventrites 3, 4 and 5 which in Scalaventer is abrupt,
glabrous, and extends the entire width of the abdomen. The abdominal modifi-
cations of some Pandeleteinus approach those of Scalaventer; however, the
abdominal modifications in these species become squamose toward the sides
and flatten out, and actually resemble the condition of Paululusus more than of
Scalaventer. Pandeleteinus has the fore coxae much more narrowly separated
and differs from the individual species of Scalaventer in two or more of the
following additional characters: beak much broader with vertical sides; ocular
vibrissae present; eyes much smaller; antenna shorter with much smaller an-
tennal club; mandibles less protruding; and corbels and dorsal surface of tarsi
squamose.
Isodrusus has the abdominal modifications similar to those of Scalaventer;
the genus differs from Scalaventer principally in its connate tarsal claws. The
West Indian Isodrusus may also be separated from Scalaventer by their pro-
portionately larger eyes and smaller antennal club; the continental Isodrusus
has ocular vibrissae.
Scalaventer is a much larger genus and much more widespread than the
other two native West Indian genera. Its 12 species occur in the Bahama Is-
lands, most of the Greater Antilles, southern Florida and Baja California; the
three species of Paululusus are known only from Hispaniola, and the two spe-
cies of Paradacrys occur in the Bahama Islands.
Biological information on the adults of Scalaventer is discussed under each
species; nothing is known of the immature stages. Figures 1 to 5 depict the
feeding damage caused by adults of various species.
Scalaventer Species Groups
I. cyrillae group. Characterized by the most extreme development of the
modification of ventrites, large and protruding eyes, narrowly separated fore
coxae, large and loose antennal club. Jamaica. Two closely related species —
which have evolved to utilize two different genera of trees for adult feeding,
Cyrilla and Coccoloba. (1) cyrillae new species, (2) coccolobae new species.
Howden: West Indian Tanymecini 2
II. litoreus group. Characterized by the broad, flat beak and robust size
and shape. Jamaica. Three closely related species with litoreus and montanus
particularly having evolved to utilize different habitats, as suggested by their
names. (3) litoreus new species, (4) montanus new species, (5) remotus new
species.
lil. subtropicus group. Characterized by widely separated fore coxae,
scarcely enlarged fore legs, small eyes, small antennal club, beak with scales
typical or near-typical to apex, flattened pronotum, relatively short and stout
aedeagus. Cuba, Grand Cayman, Jamaica, southern Florida and Baja Califor-
nia. (6) subtropicus (Fall), (7) caymani new species, (8) jamaicensis new spe-
cies. Closest to this group but with narrowly separated fore coxae and extreme-
ly long aedeagus; Bahama Islands. (9) convexifrons new species.
IV. gelinasus group. Characterized by short, thickened beak; ventrites with
concavity arcuate, much wider medially than on sides. Unique females from
Jamaica, Dominican Republic. (10) gelinasus new species, (11) valkyrius new
species.
V. cubensis group. Characterized by relatively long, narrow beak; longer,
slender legs; short aedeagus; compact antennal club. Cuba. (12) cubensis
new species.
Key to the Species of Scalaventer
1. Apical emargination of beak occupying one-third of anterior width of beak.
Head narrow between eyes; beak narrow, parallel-sided, same width as
head between eyes. Aedeagus shorter than first three ventrites measur-
eroMecrerecbeWh oleae Otol a en een eee a 12. cubensis, new species (p. 31)
Apical emargination of beak occupying one-half or more of anterior width
of, beak. Beak not narrow and parallel-sided. 2.6 ce. sen es 4 es 2
2. Ventrite 5 of female strongly, longitudinally convex its entire length and
without depressions on either side. Beak very short and thickened; dor-
sal surface strongly constricted at base where (in female) it is only one-
half as wide as head between eye (beak of male may be more slender).
Apex of beak strongly deflected so that epistoma is nearly perpendicular
to dorsal surface. High mountains of Dominican Republic. .....
ak van ad aos sc ec a ae ape wee eee 10. gelinasus, new species (p. 27)
Ventrite 5 of female flattened, or if slightly convex with a shallow depres-
Sion or pit either side near base. Beak not as above ........ 3
3. Beak broad and flat, continuous with frons, as wide as head between eyes,
epistomal suture obsolete or weak. Robust in shape and size (3.0 to 4.6
Ty Vion IRA CE cick a ee aga ae ee oo ee 2 ae ee ia 4
Beak not as above, epistomal suture usually distinctly carinate. ... 6
4, Elytra with setae erect. Blue Mountains, 4000 to 5000 feet. .....
iit 8 sda ip cic cinle Ts: ensue Acca eet ea cgelle SW a a ee 4. montanus, new species (p. 17)
Elytra with setae decumbent. Coastal or lowlands forms .......
9. Intervals 4 to 6 at their apical termination slightly produced, the swelling
conspicuous in dorsal outline. Eyes extremely flattened and large.
COC io, ele as ca ee ok ee ees 3. litoreus, new species (p. 14)
10.
i
Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
Intervals 4 to 6 not produced at their apical termination, not evident in dor-
sal view. Eyes moderately convex. Yallahs River Valley.....
Ces ote Gere ee Co ae era ae 5. remotus, new species (p. 19)
Blytral intervals Gqual, evenly Setate s.r a ed 8S 7
Elytral intervals 2, 4 and 6 less convex and/or without setae... . 8
Aedeagus as long as entire abdomen. Eye equidistant between dorsal and
ventral surfaces. Rum Cay, Bahama Islands ...........
Soe Peres Se SAGO ey es ae 9. convexifrons, new species (p. 26)
Aedeagus shorter than first three ventrites. Eye closer to dorsal surface
than to ventral surface. Southern Jamaica. .........ee-.
paces a pea Pa ee as 8. jamaicensis, new species (p. 24)
Fore coxae of both sexes separated by less than three-fourths the width of
the antennal club. Aedeagus very slender, distinctly narrower than
abeqtah Ce Se) View ie wie BAIR Wie + TE HIER y
Fore coxae of females at least separated by approximately the width or
more of the antennal club. Aedeagus (where known) as wide as antennal
Cl a ee eee FG ee ee gn ea ere ee 10
Aedeagus sinuate in profile. Male with fore femur only slightly thicker
than hindfemur. Female with glabrous anterior portion of ventrites 3,
4 and 5 grossly enlarged and modified so that the squamose surface of
ventrite 3 is perpendicular to the plane of the abdomen (Fig. 44a).
Jamaica, approximately 4000 feet andup ..........4.-.-.6
eo as a Br ee ees ee aie .% 1. cyrillae, new species (p. 7)
Aedeagus evenly arcuate in profile. Male with fore femur grossly swollen,
nearly twice as thick as hindfemur. Female with anterior portion of
ventrites 3, 4 and 5 not so grossly enlarged; the squamose portion of
ventrite 3 oblique at most, continuous with the curve of ventrite 2 (Fig.
44b). Jamaica, lowlands to approximately 4500 feet ........
PG er he en 2. coccolobae, new species (p. 10)
Scrobe acutely angled, the vertical portion slightly bowed towards apex of
beak, deep and broad to the ventral surface of the beak. Female with
anterior portion of ventrite 5 strongly arcuate medially. Blue Moun-
tains: Jamaica... 7.4 .. 11. valkyrius, new species (p. 7
Scrobe obtusely angled, not reaching the ventral surface er Rr eee
Elytral intervals 3, 5 and 7 elevated, especially towards declivity, the
elevations interrupted before declivity to form separate swellings. Su-
tural interval slightly swollen (or not) on declivity well below disc of
elytra. Cuba, Southern Florida, Baja California. .........
ge ee ge as pe eg PEN ee 6. subtropicus (Fall), (p. 21)
Elytral intervals 3, 5 and 7 scarcely more elevated than other intervals,
nowhere interrupted. Sutural interval slightly swollen at summit of
Cet yty. SPOR Cape se ek a ee ae ee es ee ee we
eee CP erate eee er er eikahce 7. caymani, new species (p. 22)
Howden: West Indian Tanymecini 7
1. Scalaventer cyrillae, new species
Figures 3, 6, 7, 16, 44a, 49, 60, 61.
Diagnosis. - Aedeagus sinuate. Female with glabrous anterior portion of
ventrite 3 enlarged, oblique instead of perpendicular, directed caudally, leav-
ing squamose portion reduced and vertical.
Description. - Holotype, male, length 2.7 mm., width0O.9 mm. Color
brown, black and pale green, marked as follows. Head mottled brown and
black; pronotum black on sides, broad median vitta predominantly pale brown.
Each elytron with a conspicuous green spot on interval 5 just before middle,
interval 4 black on basal fifth, remainder of elytron indistinctly marked. Legs
slightly annulate. Scales mostly angular; margined; reticulate on head, prono-
tum and disc of elytra, becoming granular and amarginate on sides and apex of
elytra; not contiguous; green scales of elytral spot imbricate, not margined,
finely granular. Setae completely arched, scarcely elevated, about as long as
a large scale; elytral intervals 2, 4 and 6 without setae except on declivity and
extreme base.
In profile, beak and head nearly flat from apex to posterior edge of eye
whence the contour is abruptly directed obliquely to pronotum, beak before
eyes Slightly longer than deep. Dorsum of beak at widest point slightly nar-
rower than head between eyes; edges of beak over scrobe converging towards
base, the sides here marked dorsally by a vague keel, the beak concave be-
tween these keels; median line marked only by a brief indentation between
eyes; beak deflected apicad of interantennal line which is unmarked. Apical
emargination of beak slightly obtusely angled, the sides slightly arcuate, finely
but distinctly carinate, occupying approximately half the apical edge of beak,
and with a row of three vibrissae on each side; epistoma finely rugulose.
Scales apicad of interantennal line reduced in size, number and sculpture.
Scrobe obtusely angled, dorsal edge of horizontal portion slightly sinuate,
vertical portion separated from eye and apex of beak by one scale, ending at
ventral surface beneath anterior edge of eye. Antennal club (Fig. 60) large,
‘‘loose’’, first segment cup-shaped. Eye moderate, strongly convex; viewed
anteriorly, combined depth of eyes at most three-fourths as wide as distance
between them.
Prothorax 1.05 times longer than wide, sides strongly rounded between
conspicuous, broad basal and apical constrictions. Pronotum 1.6 times longer
than prosternum; in profile, arcuate medially, the apical projection over head
conspicuously higher than disc of pronotum. Punctures deep on sides of pro-
thorax, becoming shallower medially.
Elytra 2.5 times longer than prothorax, elytra across humeri 1.2 times
wider than prothorax. Base of elytra straight. Elytra with sides very slightly
divergent from base to apical three-fifths, thence convergent, scarcely rounded,
to apex; apical terminus of intervals 4 to 6 only obsoletely interrupting outline.
In profile, elytra flat from base to about apical two-fifths, thence descending
obliquely (more strongly after apical fifth) to apex; summit of declivity broadly
rounded, poorly marked. Basal fifth of intervals 3 to 5 slightly, conjointly
convex; interval 5 obsoletely elevated just beyond middle. Elytral striae set
with fine punctures, slightly imperfectly aligned in vicinity of swellings.
Fore leg (Fig. 49) distinctly larger than hind leg; fore femur moderately,
gradually swollen, 1.3 times as thick as hind femur; fore tibia with six small
teeth on inner edge.
Fore coxae very narrowly separated, i.e., by less than the greatest width
8 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
of the scape, approximately three-sevenths the width of the antennal club.
Modification of anterior portion of ventrites 3, 4 and 5 (Fig. 16) deep, per-
pendicular, straight, the edge not carinate. Ventrite 5 slightly convex, its
apex broadly rounded and with a minute emargination.
Aedeagus (Fig. 61) in profile slightly sinuate for apical three-fifths, rather
strongly deflected for basal two-fifths, thicker at base of apical opening, apex
oblique; in dorsal view apex elongate-oval; slightly longer than first four ven-
trites.
Allotype, female, length 3.6 mm., width 1.4 mm. Differs from holotype
in the following respects. Vividly marked (Fig. 7) as follows: vertex white
with a basal black dot; pronotum with a broad, white median vitta; elytron
without a green spot; basal seventh of interval 4 black; elytral disc witha
black diamond on central half extending to stria 6 laterally, the diamond broad-
ly bordered with white anteriorly and with mixed tan and white posteriorly;
each elytron with two elongate black dots, one on interval 5 just in front of
diamond and one on interval 5 at its apical terminus; legs annulate. Scales
reticulate on head only; scales of pronotum strongly margined and coarsely
granular; scales of elytra coarsely granular and finely margined on disc, be-
coming amarginate and more finely granular toward sides and apex. Beak with
arcuate apical half nearly glabrous with only a few scattered, convex, shining
scales; median line finely carinate in this glabrous area from concavity to apex
of emargination; apical emargination approximately right-angled with sides
slightly curved. Scrobe separated from apex of beak and eye by width of two
scales. In profile, prothorax much thicker - 1.2 times longer dorsally than
thick, the pronotum 1.4 times longer than the prosternum. Elytra 3.0 times
longer than the prothorax; elytra across humeri 1.4 times wider than the pro-
thorax. Base of elytra feebly arcuate. Elytra with sides divergent from base
to just beyond middle, thence convergent, nearly straight, to the attenuate apex,
the apical terminus of intervals 4 to 6 briefly interrupting outline. In profile
(Fig. 6), the convex base of intervals 3 to 5 evident, disc subarcuate medially,
summit of declivity broadly rounded, declivity feebly concave, attenutate. Su-
tural and eleventh intervals swollen into a small knob at their apical terminus.
Elytra flatter with swellings as in male. Elytral setae as in male except be-
ginning at basal third on interval 6. Fore coxae separated by distance equal
to greatest width of antennal scape, approximately five-sevenths the width of
antennal club.
Ventrite 2 with its posterior-lateral angles enlarged, hooking over edge of
elytra. Ventrite 3 (Fig. 44a) with glabrous anterior portion greatly enlarged,
oblique instead of perpendicular, directed caudally, leaving squamose portion
reduced and vertical instead of horizontal, the lateral edges of the glabrous
keel narrowly hooking over edges of elytra. Glabrous anterior portion of
ventrites 4 and 5 likewise oblique, but only a little higher than the plane of
abdomen, the edge carinate; ventrite 4 with anterior and posterior edges
slightly arcuate, converging medially, leaving squamose portion constricted
medially; ventrite 5 elongate, approximately two-thirds as long as wide, apex
narrowly rounded, slightly convex along median line, especially basally.
Type series. - Holotype, male, Hardwar Gap, Jamaica, 4000 feet, 25
August 1966, A. T. Howden, on Cyrilla racemiflora (Howden). Allotype,
female, same data as holotype but 24 July 1966 (Howden). Paratypes, 367
males, 254 females. JAMAICA: 35 males, 22 females, same data as holo-
type (Howden); 155 males, 120 females, same data as holotype but 3-31 July
1966, A. T. Howden or [H. F.] Howden and Becker Collectors, some on
Clethra occidentalis and Alchornea latifolia (CNC, Howden); 1 male, 3 females,
Howden: West Indian Tanymecini 9
Hardwar Gap, Portland, 23 November 1958, 24 May 1963, 25 May 1964, 16
August 1964, T. H. Farr Collector (Inst. Jam.); 1 female, Hardwar Gap, 4800
feet, 13-15 July 1960, C. and P. Vaurie Collectors (AMNH); 1 male, 1 female,
Green Hills [near Hardwar Gap], Portland Parish, 21 September 1945, E. L.
Sleeper Collector (Sleeper); 1 male, Cinchona, 26 February 1911 (AMNH); 172
males, 105 females, Blue Mountain Peak, 7400 feet, 27-28 July 1966, Howden
and Becker (CNC); 1 male, Main Range, Blue Mountains, 5-7388 feet, 17-19
August 1934, P. J. Darlington (MCZ); 1 male, 1 female, Newcastle, Mile 18
[from Kingston], St. Andrew, 25 July 1966, A. T. Howden, on Cyrilla racemi-
flora (Howden).
Discussion. - Males vary in length from 2.4 to 3.4 mm. and in width from
0.7 to 1.7 mm.; females vary in length from 2.9 to 4.2 mm. and in width from
1.1 to 1.7mm. Color of males is usually similar to that of the holotype, but
one-tenth of the specimens have additional green elytral spots on interval 5 at
the apical third and on the sutural interval at the summit of the declivity; rarely
the elytral spot is white, blue or absent. Many males have the ventral surface
partly metallic golden green. Females are often vividly marked as in the allo-
type, but when the white band bordering the black diamond is broken it may be
replaced by a spot of white (one-thirteenth of the specimens) or of green (rarely)
in the same position as the elytral spot of the males. The holotype and allotype
represent the extremes of variation in head and beak except that the median line
is sometimes impressed from between the eyes to the concavity. On the disc
of the elytra, white setae especially are sometimes less strongly arched making
them much more conspicuous though they may actually be no longer. Setae
sometimes are multiserial on interval 5 and the sutural interval at the summit
of the declivity. The elytra of females particularly are variable; compared to
the allotype they may be as flat or considerably more convex and the apex may
be less attenuate. The number of teeth on the fore tibia varies from three to
seven; they are never very large. The abdomen of the female is also subject
to some variation. Blue Mountain Peak specimens do not have the ‘‘hooks’’ of
ventrites 2 and 3 as well developed and the glabrous flange of ventrite 3 is
never as enlarged as in the allotype. Hardwar Gap specimens more often have
the ‘‘hooks’’ developed to the same extent as in the allotype. The glabrous an-
terior portion of ventrites 3 and 4 is always oblique and caudally directed and
at its weakest is much more developed than in any other species of Scalaventer.
Males often have the modification of ventrites 3, 4 and 5 arcuate instead of
straight as in the holotype. The aedeagus is always strongly arcuate basally
and with some Sinuation in the apical portion.
8S. cyrillae males are unique in the sinuate aedeagus and females are unique
in the grossly modified, oblique anterior portion of ventrites 3, 4 and5. The
species may also be distinguished from its nearest relative, S. coccolobae, by
its margined, never pustulate scales; gradual elytral declivity and attenuate
apex; and less swollen fore femora. There is some overlap of the range of
cyrillae and coccolobae, but the two species appear to be host specific, at
least as adults; see Biology for further discussion.
It would seem possible that a specific barrier to mating between cyrillae
and coccolobae is evident in their morphology. The elongate fifth ventrite of
cyrillae females probably necessitates the aedeagus to be strongly arcuate at
the base for successful mating; whereas the moderate length of the fifth ven-
trite of coccolobae females will accomodate the nearly straight and shorter
aedeagus of coccolobae males.
Pandeleteius ephippiatus Champion from Panama bears a striking super-
ficial resemblance to S. cyrillae, the elytra being similarly attenuate and
10 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
marked with a white-bordered black diamond. However, ephippiatus is a true
Pandeleteius with well-developed ocular vibrissae and unmodified anterior por-
tion of ventrites 3, 4 and 5.
In this large series only two deformities were noted. In one the funicle has
segments five and six enlarged. In the other, a female, the left half of the
modification of ventrite 3 is abruptly male-like; the remainder of the abdomen
appears typically female. :
Biology. - S. cyrillae appears to be endemic to the cloud forests of the
Blue Mountain Range of Jamaica with Cyrilla racemiflora L. its preferred
adult food. In extensive tests in the summer of 1966, the weevils fed exclusive-
ly on Cyrilla leaves when offered Clethra occidentalis, Coccoloba spp., and
Cyrilla, separately or in combination. Some specimens had been collected on
Clethra but even these fed only on Cyrilla in captivity. Most of the tests lasted
three weeks, the maximum length of time the specimens and leaves could be
kept in the jars before they were completely covered with mold. Several at-
tempts to gain information on the immature stages were unproductive.
On Blue Mountain Peak, S. cyrillae was reported by H. F. Howden and
E. C. Becker to be one of the most common insects on the Peak during their
overnight stay, 27-28 July 1966. The weevils were found on Cyrilla where it
began to appear along the trail to Blue Mountain Peak and all the way to the top
of the Peak (7402 feet) even though the Cyrilla dropped out at about 6000 feet.
At the Peak the weevils occurred on Vaccinium meridionale, Clethra occiden-
talis and Clethra alexandri. As at Hardwar Gap, though, the weevils were
absent in dense growths, common on vegetation exposed to the sun.
At Hardwar Gap, the Cyrilla trees were in bloom during July and August
and the weevils were more numerous on blooming trees and new lateral shoots
off the trunk. The weevils were observed feeding on the leaves (new growth
preferred) with one long foreleg hooked over the edge of the leaf while they
gradually ate away the margin; by the end of a month many leaves were reduced
to half or less of their original size and some trees had a distinctly ragged
appearance, Figure 3 shows the typical feeding damage to Cyrilla leaves. The
weevils were as abundant on foliage during the day as at night.
The road from Kingston to Hardwar Gap offered a good opportunity to ob-
serve the junction of the ranges of S. cyrillae and S. coccolobae. Going up
from Kingston Cyrilla first appears near Newcastle at the 18 mile post; only
two S. cyrillae were taken here even though the trees were in bloom and rather
numerous. Coccoloba spp. appear as far up as the 20 mile post and a short
series of S. coccolobae was taken on these trees.
2. Scalaventer coccolobae, new species
Figures 1, 8, 13, 44b, 50, 59, 62.
Diagnosis. - Fore femur of male grossly swollen (nearly twice as thick as
the hind femur). Beak slightly constricted at the interantennal line which is
glabrous, grooved. Scales of elytral declivity pustulate.
Description. - Holotype, male, length 3.0 mm., width1.0 mm. Color
black, brown and nearly white, marked as follows. Head with a pale triangle
extending from base to frons, sides of head black. Pronotum with a pale vitta
occupying median half of disc, sides black. Elytra with intervals 1, 2 and 3
white from base to about apical third, the white area interrupted at apical third,
irregularly bordered with black; an irregular black ‘‘V’’ on apical third from
interval 5 to summit of declivity; intervals 7 and 8 entirely pale, remainder of
Howden: West Indian Tanymechini 11
elytra mottled. Legs annulate. Scales mostly angular, not contiguous except
Some pale scales; scales of dorsum of beak, vertex and apex of pronotum mar-
gined, pustulate; scales of remainder of pronotum and disc of elytra finely or
obsoletely margined, granular, not or obsoletely pustulate; most scales of
declivity pustulate, on sutural interval becoming pustulate at about middle.
Setae about as long as the longest scales; completely arched, scarcely elevated;
setae absent on intervals 2 and 4 except on declivity, very sparse on interval 6.
In profile head and beak nearly flat from apex to about middle of eye, strong-
ly convex to behind eye, thence oblique to pronotum. Dorsal surface of beak
highly sculptured; sides rather abruptly flared outwards apicad of interantennal
line where beak is slightly wider than head between eyes. Median line deeply
impressed from interantennal line to middle of eyes, sides slightly deflected
towards line. Interantennal line distinctly marked by a glabrous, shiny, slightly
V-shaped groove; apicad of line scales sparse, circular, smooth and shiny;
setae apicad of line straight, as long as two to four scales, directed towards
Sides and apex. Apical emargination ogival, occupying approximately three-
fourths of apical edge of beak, finely carinate, a short carina extending from
apex about half way to interantennal line, with a row of four vibrissae on each
Side. Scrobe slightly obtusely angled, dorsal edge of horizontal portion slight-
ly sinuate; vertical portion distinctly closer to eye than to apex of beak, sepa-
rated from eye by one scale and from apex of beak by width of about two scales;
ending at ventral surface beneath anterior edge of eye. Antennal club (Fig. 59)
large; pedunculate, i.e., first segment elongate basally. Eye large, very
strongly convex and protruding; viewed anteriorly, combined depth of eyes
approximately as great as the distance between them.
Prothorax 1.1 times longer than wide, sides moderately rounded between
moderate basal and apical constrictions. Pronotum 1.6 times longer than
prosternum; in profile, gently arcuate medially, the apical projection over
head not higher than disc of pronotum. Punctures small, shallow, inconspicu-
ous.
Elytra 2.3 times longer than prothorax, elytra across humeri 1.4 times
wider than prothorax. Base of elytra straight. Elytra with sides parallel to
apical two-thirds, thence convergent, scarcely rounded, to apex; apical ter-
minus of intervals 4 to 6 obsoletely interrupting outline. In profile, elytra
flat to apical two-thirds; summit of declivity broadly rounded; the declivity
oblique, straight. Intervals 3 and 4 obsoletely, conjointly elevated on basal
fifth; intervals 5 and 7 obsoletely elevated on apical half of disc; sutural inter-
val slightly produced at summit of declivity. Striae set with small punctures.
Fore femur (Fig. 50) grossly swollen, 1.8 times thicker than hind femur;
fore tibia with six small teeth and some serrulations on inner edge.
Fore coxae separated by less than the greatest width of the antennal scape;
separated by approximately one-third the width of the antennal club. Modifica-
tion of anterior portion of ventrites 3, 4 and 5 slightly wider medially, be-
coming nearly obsolete laterally on ventrite 3 and 4, the edge not carinate.
Ventrite 5 nearly flat, broad, the apex narrowly truncate.
Aedeagus (Fig. 62) longer than first 3 ventrites; approximately one-fifth
thicker than thickest part of scape; in profile, gently, almost evenly arcuate,
the apex strongly oblique; in dorsal view, apex attenuate-elliptical; base of
opening coriaceous along median line nearly to middle of aedeagus.
Allotype, female, length 3.5 mm., width 1.3 mm. Slightly teneral, hence
color paler. Differs from holotype in the following respects. White markings
of head and pronotum less distinct, somewhat mottled; head with a white spot
between eyes embracing end of median groove; markings of elytra reduced to
12 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
the black spots, each of which is bordered posteriorly by a white spot. Beak in
profile slightly concave; groove along median line reduced to an interocular pit
and deep triangular pit at interantennal line. Scrobe with vertical portion nar-
rower, equidistant between eye and apex of beak. Eye not as protruding, com-
bined depth of eyes approximately three-fifths as great as distance between them.
Prothorax 1.2 times longer than wide, sides straight between constrictions.
Elytra 2.5 times longer than prothorax. Sides of elytra sub-parallel to apical
third, thence convergent, slightly rounded to apex, the apex scarcely exceeding
the summit of declivity viewed dorsally. Elytra in profile (Fig. 8) with convex
base of intervals 3 and 4 evident, disc subarcuate, summit of declivity rounded,
declivity slightly oblique. Fore femur less swollen, 1.3 times wider than hind
femur; one tooth of fore tibia bifid. Fore coxae separated by approximately
two-thirds the width of the antennal club. Modification of anterior portion of
ventrites 3, 4 and 5 (Figs. 13, 44b) slightly oblique instead of perpendicular,
rounded at the bottom, slightly arcuate in ventrites 3 and 4, median half ele-
vated and subcarinate, squamose portion of ventrites 3 and 4 obsoletely con-
cave. Ventrite 5 approximately one-half as long as wide, flattened and slightly
turned up at apex, apex rounded.
Type series. - Holotype, male, Mandeville, Manchester Parish, Jamaica,
16 August 1966, A. T. Howden, on Coccoloba schwarzi (Howden). Allotype,
female, same data as holotype (Howden). Paratypes, 141 males, 106 females.
JAMAICA (listed by parishes from West to East): 55 males, 39 females, same
data as holotype, A. T. Howden or [H. F.]| Howden and Becker Collectors
(CNC, Howden); 18 males, 15 females, Barbecue Bottom, Trelawny, 4, 10,
12 August 1966, A. T. or H. F. Howden, on Coccoloba troyana and Coccoloba
longifolia (CNC, Howden); 1 male, 2 females, Good Hope, Trelawny, 11, 17
August 1966, A. T. Howden, on Coccoloba schwarzi (Howden); 5 males, 1 fe-
male, Hermitage, St. Elizabeth, 17 August 1966, E. C. Becker, on Coccoloba
schwarzi (CNC); 1 female, Hermitage Res., St. Andrew, 21 September 1945,
E. L. Sleeper (Sleeper); 5 males, 4 females, Irishtown, St. Andrew, 28
August 1966, A. T. Howden, on Coccoloba longifolia (Howden); 24 males, 20
females, Mahogany Vale, St. Andrew, 12, 20, 28 July 1966, A. T. Howden or
[H. F.] Howden and Becker, on Coccoloba tenuifolia (CNC, Howden); 1 female,
Mandeville, Manchester, 28 September 1945, E. L. Sleeper (Sleeper); 1 female,
Mandeville, A. E. Wight (MCZ); 18 males, 17 females, Mizpah, Manchester,
5, 16 August 1966, A. T. or H. F. Howden, on Coccoloba schwarzi and Coc-
coloba longifolia (CNC, Howden); 6 males, 1 female, Newcastle, St. Andrew,
25 August 1966, on Coccoloba sp., A. T. Howden (Howden); 8 males, 3 females,
Whitfield Hall, St. Thomas, 27 July 1966, A. T. Howden, on Coccoloba tenui-
folia (Howden); 1 male, 1 female, Whitfield Hall, Blue Mountains, near 4500
feet, 13-20 August 1934, Darlington (MCZ).
Discussion. - Variation in the type series is as follows. Males vary in
length from 2.4 to 3.4 mm. and in width from 0.8 to 1.3 mm.; females vary
in length from 2.7 to 3.9 mm. and in width from 1.1to1.5 mm. Males
usually have the disc of prothorax and elytra broadly white, broadly bordered
with black on the prothorax; females usually have the disc of prothorax pre-
dominantly pale but bordered with mottled black, and the disc of the elytra
rarely broadly pale. The number of pustulate scales is highly variable; the
holotype represents nearly the maximum area covered by pustulate scales;
rarely there are no distinctly pustulate scales even on the apex of the declivity.
The interantennal line is always sharply defined by a groove and usually is
conspicuously glabrous and shiny; occasionally, however, one or more scales
may partially obliterate the groove. The sides of the apical emargination of
Howden: West Indian Tanymechini 13
the beak are never straight; they may be arcuate or nearly parallel before
converging to the apex. There are four or five vibrissae on each side of the
apical emargination. In profile, the frons is sometimes much more abruptly
transversely convex than in either the holotype or allotype. The eye varies in
the distance it protrudes; in males it may be considerably less protruding than
in the holotype but always more than in the allotype and in one extreme male
the combined depth of the eyes is 1.3 times wider than the head between the
eyes. The antennal club is rather variable, being sometimes much broader
and often less pedunculate than in the holotype and allotype; however, none
were seen that were as broad at the base as in cyrillae. In a few specimens
the anterior projection of the pronotum is higher than the plane of the disc,
similar to the condition of cyrillae. Elytra sometimes have interval 3 obsolete-
ly elevated. In one striking male from Mahogany Vale, the setae on the disc
of the elytra were ‘‘uncurled’’, i.e., Semi-erect and incompletely arched.
One-fifth of the paratypes from Mahogany Vale have some elytral setae partially
uncurled, a much higher incidence than in the other localities. Many speci-
mens have the numerous setae of the summit of the declivity less arcuate and
conspicuous. The fore femora vary in the extent of their swelling; in males
they are sometimes less swollen than in the holotype and in at least one male
are swollen fully twice as wide as the hind femora. One particularly robust
female from Barbecue Bottom has the fore femora 1.6 times the width of the
hind femora. Fore tibial teeth vary from three to eight in number and are
irregular in size and placement. Lateral edges of ventrites 2 and 3 were never
observed to have hooks as in cyrillae, but a few females have an obsolete ex-
tension here. No appreciable variation was observed in the aedeagus.
Scalaventer coccolobae may always be separated from cyrillae by its gently
arcuate aedeagus and relatively simple female abdominal modifications. Other
differences are the lack of metallic green scales on ventrum and elytra, pustu-
late scales on the elytral declivity, larger eye, narrower base of antennal club,
less attenuate apex of elytra, and frons more strongly and more anteriorly
Swollen. From other Scalaventer, coccolobae may be distinguished by its
narrowly separated fore coxae, well-marked interantennal line, and grossly
swollen fore femur.
The ranges of cyrillae and coccolobae overlap slightly, but there is no evi-
dence of character displacement. S. cyrillae adults were never collected on
Coccoloba and S. coccolobae was never collected on Cyrilla; in captivity neither
species of Scalaventer would feed on the adult food of the other species.
Teratological manifestation includes two specimens with a deformed abdo-
men, one with a twisted body and one with a bifid antennal club.
Biology. - S. coccolobae is an endemic Jamaican species, widespread over
the island where it is found at elevations ranging from 500 to 4500 feet. It has
been collected on four species of Coccoloba only: longifolia Fisch., tenuifolia
L., troyana Urban, and schwarzi Meisner.
The Mandeville series was collected in an open hillside pasture with
scattered, apparently mature, C. schwarzi. Each tree seemed heavily infested
and beating yielded numerous weevils (as well as a shower of ants). The Mizpah
series was taken on a large tree in full bloom and most of the specimens seemed
to come from the vicinity of the flower racemes. Most of the other specimens
were taken on saplings growing along the roadside, sometimes in dense growths.
Not all Coccoloba trees yielded Scalaventer and the reason for some being very
productive while others in apparently similar situations were barren was never
discovered.
Figure 1 shows the typical feeding damage of adult S. coccolobae on the
14 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
leaves of C. tenuifolia. Note that the weevils fed on the interior of the leaf as
well as from the margin inwards. This appears to be a characteristic of the
species of Scalaventer rather than an effect of a more tender host leaf. When
Scalaventer cyrillae fed on their favorite young leaves it was always exclusive-
ly along the margin.
3. Scalaventer litoreus, new species
Figures 4, 5, 14, 38, 48, 63.
Diagnosis. - Beak flat. Eyes very large, strongly flattened. Punctures
on sides of pronotum deep to foveate. Elytral setae strongly curved, their
apices not quite touching surface.
Description. - Holotype, male, length 3.4 mm., width 1.3 mm. Color
dark brown, tan and white, indistinctly marked. Head mottled, with a dark
brown vitta behind eyes and a dark brown triangle on occiput; pronotum in-
distinctly marked with a pair of pale parentheses-shaped vittae on disc, sides
dark brown; each elytron with a whitish spot on intervals 5 and 6 just before
middle, apex pale beyond a ‘‘V’’ on apical third bordered anteriorly with dark
brown; tibiae and to a lesser extent femora annulate with brown and white.
Scales of irregular rounded or angular shapes, moderately to coarsely granu-
lar, margined on head and pronotum, weakly margined on elytra, not contigu-
ous. Setae of head and pronotum nearly prostrate, as long as one to two
scales; setae of elytra (Fig. 48) broader, evenly strongly arcuate, their apices
not quite touching the surface, uniserial on intervals, directed apically except
on sutural interval at summit of declivity where three setae are directed medi-
ally.
In profile, beak obsoletely arcuate to behind middle of eyes, thence gradu-
ally rounded to pronotum. Dorsum of beak as wide as head between eyes,
rounded on edges, median line impressed for a short distance at base of frons.
Interantennal line unmarked. Apical emargination approximately right-angled,
finely but distinctly carinate, occupying approximately three-fifths of apical
edge of beak, with a row of five vibrissae on each side; surface of epistoma
finely rugulose. Beak slightly depressed behind apical emargination, glabrous
for about the width of one scale; with only three or four rows of scales simpli-
fied before they become typical in size and sculpture. Scrobe (Fig. 38) very
obtusely angled; horizontal portion brief, slightly arcuate; vertical portion
oblique, separated from eye and apex of beak by two scales, ending at ventral
surface beneath anterior edge of eye. Antennal club elongate, elliptical. Eye
very large, subcircular, only slightly convex.
Prothorax as long as wide, sides gently rounded between narrow basal and
apical constrictions. Pronotum 1.6 times longer than prosternum; pronotum
in profile flattened, 1.2 times longer dorsally than thick, disc scarcely arcu-
ate between constrictions. Punctures of prothorax foveate on sides, rapidly
becoming obsolete on disc.
Elytra 2.5 times longer than prothorax, elytra across humeri 1.3 times
wider than prothorax. Base of elytra obsoletely arcuate. Sides of elytra
very slightly divergent to about middle, thence gently converging to apical
sixth where the apical terminus of intervals 4 to 6 is prominent in the dorsal
outline as an obliquely angled protuberance, the apex rounded beyond this. In
profile elytra flattened from base to about middle, thence slightly descending
to summit of declivity which is broadly rounded (Fig. 38). Elytral striae
marked with large, deep punctures on basal half, the punctures becoming much
Howden: West Indian Tanymechini 5
smaller beyond middle; elytral intervals slightly convex, about as wide as the
distance between punctures in striae.
Fore femur rather strongly, abruptly swollen medially; fore tibia with four
very small teeth on inner edge of one leg, three teeth on other leg.
F ore coxae separated by distance equal to about two-thirds the width of the
antennal club. Anterior portion of ventrites 3, 4 and 5 abruptly, perpendicular-
ly concave; the edge not carinate, on ventrite 5 slightly arcuate medially. Ven-
trite 5 convex, apex broadly rounded.
Aedeagus arcuate (Fig. 63); as long as first four and a half ventrites; api-
cal opening elongate-elliptical, in profile more oblique than in montanus.
Allotype, female, length 4.6 mm., width 1.8 mm. Specimen slightly tener-
al, coloring less distinct than in male. Differs from holotype in the following
respects. Sutural interval at summit of declivity with five conspicuous, white,
medially directed setae; apical terminus of intervals 4 to 6 with two conspicu-
ous, apically-directed setae. Head and beak much more robust than in type,
dorsum of beak slightly narrower than head between eyes. Beak glabrous
medially from apex of apical emargination to interantennal line, the glabrous
area finely rugulose as are the epistoma and dorsum of mandible. Punctures
of prothorax and elytra shallower than in holotype. Elytra in dorsal view with
sides more rounded, apex a little more attenuate and apical terminus of inter-
vals 4 to 6 more prominent. In profile elytra with declivity obsoletely concave,
marked at its summit by a slight swelling. Inner edge of fore tibia with four
teeth on one side, five on the other.
Anterior portion of ventrites 3, 4 and 5 (Fig. 14) abruptly, perpendicularly
concave, the edge subcarinate on ventrites 4 and 5, central three-fifths rather
strongly arcuate, and on ventrites 4 and 5 elevated medially as well. Ventrite
5 nearly flat, its apex narrowly rounded. |
Type series. - Holotype, male, Duncans, Trelawny, Jamaica, 19 August
1966, A. T. Howden, on Krugiodendron ferreum (Howden). Allotype, female,
same data as holotype (Howden). Paratypes, 143 males, 117 females. JAMAI-
CA: 79 males, 53 females, same data as holotype except 28 on Krugiodendron
ferreum, 47 on Haematoxylum campechianum, A. T. Howden or [H. F, How-
den and Becker Collectors (CNC, Howden); 59 males, 57 females, same locali-
ty as holotype, 8-23 August 1966, A. T. Howden or [H. F.]| Howden and Becker,
2 at black light, 54 on Krugiodendron ferreum, 73 on Haematoxylum campechi-
anum, 3 on Ehretia tinifolia (CNC, Howden); 5 males, 5 females, Reading, St.
James, 18 August 1966, A. T. Howden, on Haematoxylum campechianum
(Howden); 2 females, Alligator Pond Bay, 20 February 1937, Chapin and Black-
welder, Sta. 410 (USNM). |
Discussion. - Males vary in length from 3.0 to 4.1 mm. and in width from
1.2 to 1.6 mm. ; females from the north coast (Duncans, Reading) vary in
length from 3.4 to 4.6 mm. and in width from 1.4 to 2.0 mm. The two para-
types from the south coast (Alligator Pond Bay) are particularly robust females
0.4 to 5.5 mm. in length and 2.2 to 2.3 mm. in width, fully one-sixth larger
than any north coast females. These two specimens also differ from the north
coast specimens in having the scrobe scarcely angled, the elytral setae more
numerous, and the sutural interval much more produced at the summit of the
declivity, in the latter aspect approaching the condition of remotus. Variation
in color pattern is considerable, the brown and white bands on the femora and
tibiae being the most constant. The thorax is seldom as well marked as in the
holotype and allotype. The elytral spot on intervals 5 and 6 is often complete-
ly absent, is not associated with sex, and in a very few specimens is greenish.
Maximum color pattern is found in females which have the elytral white ‘‘V’’
16 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
particularly sharp and broad, often covering the entire declivity but with the
apical terminus of intervals 4 to 6 darker. Some features of the beak are sub-
ject to considerable variation. The apical emargination is usually approxi-
mately right-angled, but may be acutely or obtuSely angled; the sides of the
apical emargination may be straight or curved and they may be a little more
strongly carinate than in the holotype. The holotype and allotype represent
the two extremes of the condition of the scales on the apex of the beak. The
presence of the interocular pit is quite constant, though it is generally deeper
in males than in females and in males is usually deeper than in the holotype;
it may be reduced to a finely impressed line. The scrobe varies in length
from complete to the ventral surface to ending well above it, depending partly
on the robustness of the beak.
The elytral declivity of males is often feebly sinuate and sometimes has the
summit feebly indicated. Compared to the allotype, the elytra of females may
have the declivity more or less concave, the sutural interval much more pro-
duced at the summit of the declivity, and the apex a little more attenuated.
The females often have the modification of the anterior portion of the ventrites
less arcuate than in the allotype, but the modification of ventrite 5 at least is
always finely carinate and slightly elevated medially. Ventrite 5 of females is
usually more convex along the median line than in the allotype.
S. litoreus is very closely related to montanus (see discussion of montanus),
the strongly curved elytral setae of litoreus being the most reliable physical
character for separating the two species. If specimens of both species are
available for comparison, it is obvious that litoreus also has a more strongly
carinate epistoma, paler color, more shallowly punctured pronotum, and more
obtusely-angled scrobe. On the average, litoreus also has a flatter eye and
more abrupt declivity, but there is overlap on these characters. S. litoreus is
known only from small areas on the north and south coasts of Jamaica, where-
as montanus occurs in the Blue Mountains between about 4000 and 5000 feet.
From other species of Scalaventer, litoreus may be separated by its very
flat beak; large, flat eyes; terminus of intervals 4 to 6 prominent in dorsal
view.
Teratological manifestations include two deformed abdomens, one deform-
ed beak, and a bifid tibia with one normal tarsus and one bifid tarsus.
Biology. - At Duncans, where the majority of the type series was collected,
the first few specimens were taken in the morning on Krugiodendron ferreum
(Vahl) Urban, Ehretia tinifolia L., and Haemotoxylum campechianum L. (log-
wood). Here the three species of trees were scattered in a fence row and ina
hilly cow pasture overgrown with Opuntia. The beetles were never found in
numbers on these trees during the day. At night the Krugiodendron and Ehretia
still yielded only sparse numbers of beetles, but several large logwood trees
on an exposed slope were found to be heavily infested. The beetles were very
active at night and when a large number were beaten onto a collecting sheet
some would always escape the collector by flying off. This is quite in contrast
to other Scalaventer (except montanus) which, like Pandeleteius, land ‘“‘ spread
eagle’’ on the sheet and remain motionless for a long time.
S. litoreus was taken at only one other place on the north coast - on logwood
growing in an undeveloped residential lot at Reading, St. James. Logwood was
beaten extensively elsewhere and yielded no Scalaventer.
In feeding experiments, specimens were kept segregated according to the
tree on which they had been collected; each jar was provided with leaves of all
three species of tree. All specimens fed on Krugiodendron ferreum extensive-
ly and to a much lesser extent on logwood. There was no evidence of feeding on
Howden: West Indian Tanymechini 17
Ehretia tinifolia after a week. The typical feeding damage of S. litoreus on
leaves of Krugiodendron is shown in Fig. 4 and on leaves of logwood in Fig. 5.
Note that Krugiodendron leaves especially were eaten from the margin inwards
in contrast to the feeding of other species except S. coccolobae. These Krugio-
dendron leaves were relatively thin and tender, and the Coccoloba leaves near-
ly as tender.
A possible explanation for the distribution of S. litoreus may be found in
the adult food. Krugiodendron, native to Jamaica, occurs in dry limestone
woodland; it is not known from east of Clarendon or St. Ann Parishes or from
the extreme western part of the island. Logwood occurs commonly in dry
thickets at low elevations all over Jamaica, but it is not native.
4. Scalaventer montanus, new species
Figures 2, 10, 17, 65.
Diagnosis. - Elytral setae erect. Thoracic punctures large or foveate on
Sides.
Description. - Holotype, male, length 3.5 mm., width 1.3 mm. Specimen
slightly teneral as evidenced by the testaceous color of the aedeagus and the
submetallic lustre of the dorsal scales; abdomen slightly dislocated. Color
dark and light brown, indistinctly marked. Head with a dark basal triangle;
pronotum with an imperfect elongate, dark, median diamond enclosed by pale
‘‘narentheses’’; elytra with a vague, pale ‘“‘V’’ on apical third, and an irregular
cluster of pale green scales on intervals 5 and 6 before middle; tibiae with a
narrow pale band around the middle. Scales of irregular angular shapes,
coarsely granular, mostly margined, not contiguous. Setae of head and prono-
tum inconspicuous, nearly prostrate, slender, as long as two to three scales.
Setae of disc of elytra slightly longer, lanceolate, erect for approximately
basal two-thirds with the apical third deflected; uniserial with some irregulari-
ties; directed apically except on sutural interval at summit of declivity where
four are directed medially.
In profile, beak obsoletely arcuate to middle of eyes, thence arcuate to
pronotum. Dorsum of beak very slightly narrower than head between eyes,
the edges broadly rounded from eyes to antennal insertions. Median line finely
impressed from interantennal line to beyond middle of eyes, ending in a narrow
pit. Interantennal line unmarked. Beak apicad of interantennal line remarkably
flat; scales abruptly very small and sparse; apical emargination not well de-
fined, obtusely angled, occupying approximately three-fifths of apical edge of
beak, with a row of four slender vibrissae on each side; surface of epistoma
finely rugulose. Scrobe obtusely angled with horizontal portion very short,
arcuate; widened at angle; vertical portion oblique, deep throughout, ending at
ventral surface beneath anterior edge of eye, at closest points separated from
apex and eye by about width of two scales. Antennal club elongate, elliptical.
Eye large, nearly circular, moderately convex.
Prothorax as long as wide, sides gently rounded between narrow basal and
apical constrictions. Pronotum 1.7 times longer than prosternum; pronotum in
profile flattened, disc slightly arcuate between constrictions, 1.3 times longer
dorsally than thick. Punctures of prothorax foveate on sides, becoming shal-
lower medially.
Elytra 2.6 times longer than prothorax, elytra across humeri 1.2 times
wider than prothorax. Base of elytra slightly arcuately emarginate, sides of
elytra very slightly divergent from base to about middle, thence gently converg-
18 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
ing to apex, the apical terminus of intervals 4 to 6 prominent in dorsal outline
as an obliquely angled protuberance. In lateral view (Fig. 10), elytra obsolete-
ly arcuate from base to about middle, thence gradually descending to apex, the
summit of declivity scarcely perceptible. Elytral striae 1 to 9 marked with
large, deep punctures, the punctures becoming smaller beyond middle; elytral
intervals slightly convex, basally about as wide as space between punctures in
striae.
Fore femur rather strongly, abruptly swollen medially; fore tibia with five
small, equidistant teeth on its inner edge, serrulate apicad of last tooth.
Fore coxae separated by distance equal to three-sevenths the width of the
antennal club. Anterior portion of ventrites 3, 4 and 5 abruptly, perpendicular-
ly concave, the edge not carinate, slightly arcuate medially. Ventrite 5 con-
vex, apex rounded.
Aedeagus (Fig. 65) arcuate, as long as first four and one-third ventrites;
apical opening elliptical.
Allotype, female, length 4.6 mm., width 1.8 mm. Differs from holotype
in the following respects. Vividly marked as follows: pronotum mostly pale,
the black median diamond reduced to a few fine black lines; elytra with a con-
Spicuous white vitta extending along interval 7 for basal third, along intervals
6 and 7 for median third, thence obliquely to suture at summit of declivity,
the disc of elytra broadly black in front of the white, remainder of disc mottled
brown; tibiae with central band conspicuous, an apical band less so; the cluster
of green scales on intervals 5 and 6 greatly reduced, inconspicuous. Setae of
elytra the same color as the scales beneath them. Head much more robust,
eyes flatter. Pronotum with punctures shallower. Proportions of pronotum
and elytra as in holotype. Elytra with strial punctures much smaller than in
holotype, moderate, becoming small apically. Sutural interval moderately
produced at summit of declivity; intervals 3 and 5 convex at summit of declivity
only, apical terminus of intervals 4 to 6 slightly more produced than in holo-
type, all these swellings with a few extra setae. Fore tibia with only three
equidistant teeth on inner edge. Anterior portion of ventrites 3, 4 and 5 (Fig.
17) with the modification more distinctly arcuate medially, the edge subcarin-
ate in ventrite 4, carinate in ventrite 5. Ventrite 5 scarcely more elongate
than in holotype, obsoletely convex medially.
Type series. - Holotype, male, Whitfield Hall, St. Thomas, Jamaica, 27
July 1966, A. T. Howden, on Clethra occidentalis, Feed. Exp. [= used in feed-
ing experiments] (Howden). Allotype, female, same data as holotype (Howden).
Paratypes, 13 males, 5females. JAMAICA: 8 males, 4 females, same data
as holotype (Howden); 1 female, Whitfield Hall, Blue Mountains, near 4500 feet,
13-20 August 1934, Darlington (MCZ); 1 male, Hardwar Gap, 4000 feet, 9 July
1966, [H. F.] Howden and Becker (CNC); 3 males, Hardwar Gap, 4000 feet, 10,
15 July 1966, A. T. Howden, one specimen collected at light (Howden); 1 male,
Cinchona to Newhaven Gap, St. Andrew, 27 August 1949, Robert Hart (Sleeper).
Discussion. - Males vary in length from 3.3 to 3.8 mm. and in width from
1.3 to 1.4 mm.; females vary in length from 4.0 to 4.5 mm. and in width from
1.6to 1.8 mm. The green elytral spot is not as uniquely masculine as it is
in cyrillae; of the montanus paratypes, five males have no green spot and two
females do have some green. The holotype represents the usual coloring of
males; one male has the white elytral line developed almost as much as in the
allotype. The allotype is by far the most vividly marked specimen. The setae
may be all pale or the same color as the scales beneath. The interocular pit
on the median line is absent in one male and is often greatly reduced or absent
infemales. The poorly defined epistoma is remarkable; it is less distinct than
Howden: West Indian Tanymechini 19
in Hadromeropsis, but more so than in Tanymecus. The teeth of the fore
tibiae vary in number from three to five, often on different sides of the same
beetle. The fore coxae are much closer in most males than in the holotype.
The aedeagus varies from moderately to strongly arcuate in profile; the figure
represents the median.
S. montanus is very close to litoreus, the erect elytral setae of montanus
being the only tangible character by which one can reliably separate the two
Species. However, the species appear to occupy two distinctly different habi-
tats: cool mountain cloud forest vs. hot sea-coast; montanus is known only
from about 4000 to 5000 feet in the Blue Mountains where Clethra occidentalis
is the preferred adult food, and litoreus is known only from sea level where
Krugiodendron ferreum and Haematoxylum campechianum are the preferred
adult foods. In comparison with litoreus, montanus tends to have a less dis-
tinct epistoma, darker color (quite in keeping with its geographical habitat),
more deeply punctured pronotum, and more angled scrobe. The erect elytral
setae will also distinguish montanus from all other Scalaventer.
At Hardwar Gap, montanus occurs sympatrically with cyrillae, and at
Whitfield Hall montanus occurs with coccolobae. These species belong to the
species group characterized by a slender, highly sculptured beak, grossly pro-
truding eyes, and highly modified female ventrites.
Biology. - Clethra occidentalis appears to be the usual food of montanus
adults, although specimens kept in jars for two weeks with both Clethra and
Coccoloba leaves, fed on the Coccoloba leaves as well. They were not tested
for Cyrilla feeding. At Whitfield Hall, montanus was collected only on Clethra
and S. coccolobae was the only species collected on Coccoloba. At Hardwar
Gap where both Clethra and Cyrilla occur but not Coccoloba, three specimens
of montanus were collected, one at night at black light and two without host
data. A few specimens of S. cyrillae were taken on Clethra but in extended
feeding experiments they did not feed on Clethra whether given a choice of
leaves or not.
Figure 2 illustrates the typical feeding damage of montanus to Clethra
leaves: margins excised continuously with little feeding inwards and no feeding
on stems or petioles.
The population of montanus is certainly much larger at Whitfield Hall, where
an overnight trip yielded 14 specimens, than at Hardwar Gap where several
weeks of intensive collecting yielded only three specimens. This coincides with
the abundance of Clethra. The season of adult emergence is indicated by the
fact that teneral specimens were collected at both ends of the period 9 July to
27 August. The Howden series from Whitfield Hall was taken along the first
mile of the trail leading to Blue Mountain Peak where Clethra trees of good
size were just coming into bloom. The specimens were all beaten from the
foliage of the trees and none were observed prior to beating.
9. Scalaventer remotus, new species
Figures 20, 39, 43, 47.
Diagnosis (based on female). - Elytra with sutural interval at summit of
declivity greatly elongated into a conical protuberance. Carina of anterior
portion of ventrite 5 abruptly produced into a point medially. Pronotum with
punctures small to obsolete. Elytral setae scarcely arched, their apices touch-
ing surface.
Description. - Holotype, female, length 3.4 mm., width 1.4 mm. Color
20 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
rust, gray-white and dark brown-black. Head obscure rusty with dark vitta
behind eye, side of head and beak whitish; pronotum predominantly gray-white
with an irregular dark vitta at extreme lateral-dorsal edge. Elytra elaborately
patterned as follows: disc rust except sutural and second intervals gray, be-
coming darker posteriorly; humerus and interval 4 on basal eighth dark; a
dark, sinuate fascia across basal third from suture to interval 7 partly en-
closing a white spot on intervals 5 and 6; a dark lunule on apical third from
suture to stria 7; a broad, conspicuous white lunule just behind dark lunule,
this white lunule becoming tan posteriorly and with a short dark vitta on inter-
vals 3, 5 and 7 on or before declivity; sides of elytra medium gray. Femora
obsoletely banded; ventral surface subopalescent gray-white. Scales of irregu-
lar angular shapes; scales of head and pronotum mostly contiguous; granular to
subreticulate, not margined or very weakly so; scales of elytra mostly not
quite contiguous, not margined but some scales with a thickening of the granules
around the edge. Setae of head and prothorax as long as the longer scales,
nearly prostrate, moderately thick, dark on head and apex of pronotum, white
on remainder of prothorax. Setae of elytra (Fig. 47) like those of head and
pronotum but slightly more arched, their apices still touching surface; setae
same color as the scales beneath them, uniserial on all intervals but more
numerous on alternate intervals and on sutural protuberance.
Beak (Fig. 43) short, thick, cubical; in profile (Fig. 39), beak flat, ina
continuous plane with frons, head gently arcuate from posterior edge of eyes.
Dorsum of beak as wide as head between eyes, feebly excavate between in-
sertion of antennae. Interantennal line unmarked. Median line impressed for
a short distance between eyes; beak with a vague, shallow depression at base,
the depression splitting and continuing briefly around apical emargination;
apical emargination ogival, occupying one-half the apical width of the beak,
weakly carinate posteriorly, a weak glabrous keel continuing from apex to
depression at base of beak. Scales apicad of interantennal line small, convex,
slightly granular; setae apicad of interantennal line rapidly becoming longer,
more erect, and more numerous apically, with a row of eight to eleven vibris-
sae on each side of apical emargination. Scrobe obtusely angled, deep through-
out its length; dorsal edge of horizontal portion sinuate, wider at angle; verti-
cal portion with its posterior edge slightly curved, at the narrowest point
equidistant between apex and eye from which it is separated by only one scale,
ending at ventral surface beneath about middle of eye. Antenna with first seg-
ment of funicle elongate-oval; second segment half the size of the first; seg-
ments three to seven shorter, equal in length, cubical, becoming broader than
long; antennal club moderate with all four segments discrete. Eye moderate,
elongate.
Prothorax as wide as long, wider apically than basally, sides moderately
rounded between broad apical and basal constrictions, the constrictions nearly
confluent on disc medially. Pronotum 1.7 times longer than prosternum; pro-
notum sinuate in profile, 1.2 times longer dorsally than thick. Prothorax with
punctures small to obsolete on disc and sides, except in the dark lateral vitta
where they are small to moderate.
Elytra 2.7 times longer than prothorax, elytra across humeri 1.4 times
wider than prothorax. Base of elytra slightly arcuately emarginate. Elytra
in dorsal view with sides slightly divergent from just behind humeri to about
middle, thence gently rounded to apex, slightly constricted beneath apical
terminus of intervals 4 to 6. Elytra in profile (Fig. 39) with basal sixth
somewhat prominent, thence slightly arcuate to short, conical protuberance
on sutural interval at summit of declivity, dorsal surface of protuberance
Howden: West Indian Tanymechini 21
nearly continuous with the curve of the elytra, protuberance set about one-
third below disc of elytra; the declivity nearly perpendicular, slightly concave,
its apex slightly exceeding the end of the protuberances. Elytral striae set
with small, perfectly aligned punctures; elytral intervals slightly convex, api-
cal terminus of intervals 4 to 6 not produced.
Fore femur gradually, moderately swollen medially; fore tibia scarcely
expanded apically, with four small, equidistant teeth on inner edge.
Fore coxae separated by a distance equal to width of antennal club. Anteri-
or portion of ventrites 3, 4 and 5 (Fig. 20) abruptly concave, the modification
slightly arcuate; ventrite 4 with a median tubercle, ventrite 5 with median tuber-
cle expanded into an abrupt median point from which a short keel extends caudal-
ly. Ventrite 5 flat except for median point, elongate.
Type series. - Holotype, female, Mahogany Vale, St. Andrew, Jamaica,
20 July 1966, A. T. Howden, on Coccoloba tenuifolia (Howden). No paratypes.
Discussion. - It is unfortunate that the two prime diagnostic characters
of this species are almost uniquely female. Undoubtedly males will have no
such projected point on ventrite 5, and the conical protuberance of the sutural
interval will be considerably reduced. Unfortunately, the distinctive color and
pattern of the female may also be completely different in the male, judging from
congeners. The male, of course, may well have a distinctive aedeagus, if not
other characters of its own. In any event, the scrobe is seldom subject to sexu-
al modification and it, combined with the characters of the beak should separate
either sex of remotus from other Scalaventer species.
From other Scalaventer in the species group (montanus, litoreus) with a
broad, flat beak, remotus may be separated by its more numerous vibrissae
bordering the apical emargination, obsolete punctures of pronotum, nearly
prostrate elytral setae, and unmodified apical terminus of intervals 4 to 6.
Biology. - The holotype was collected about midday on one of three trips
to Mahogany Vale during July, 1966. The locality is in the interior Yallahs
River Valley at a relatively broad flood plain with a scrub growth at the edge
of the plain containing sapling Coccoloba tenuifolia L. Numerous specimens
of S. coccolobae were taken at the same time and it is possible that night
beating would have produced more specimens of both species. It was extremely
hot during the day.
6. Scalaventer subtropicus (Fall), new combination
Pandeletejus [sic] subtropicus Fall, 1907, p. 263; Howden, 1959, p. 387-9.
Lectotype, here designated, male, labelled, ‘‘Key Largo, Fla.’’ [hand
printed], ‘‘subtropicus [hand printed] Type [mechanically printed on
white paper]’’, ‘‘“MCZ Type 25140’’ [on red paper] (MCZ).
Diagnosis. - Elytral intervals 3, 5 and 7 elevated, especially towards the
declivity, the elevations interrupted before declivity to form separate swellings.
Sutural interval slightly swollen (or not at all) on declivity well below disc of
elytra.
Description. - Since this species was recently discussed in detail (Howden,
1959), only a few pertinent details of comparative value are recorded here.
Length 2.9 to 4.0 mm. Beak fully squamose with typical or nearly typical
scales to apex; apical emargination with an average of two vibrissae on each
side. Beak with median line impressed from interantennal line to middle of
eyes or beyond; interantennal line marked with a faint, fully squamose ridge;
22 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
median line marked or not with a faint glabrous carina from apex of epistoma
to interantennal line.
Sides of elytra in both sexes gently divergent to about middle thence gradual-
ly converging to apex, the apical terminus of intervals 5 to 7 swollen and inter-
rupting dorsal outline. All elytral swellings more prominent in females than in
males. Declivity gradual, not marked at its summit. Elytra with strial punc-
tures slightly irregularly aligned on disc.
Females with anterior portion of ventrites 3, 4 and 5 with concavity per-
pendicular, sharp-edged but not carinate, nearly straight; ventrite 5 nearly
flat with a distinct shallow pit at base either side of midline. Males with con-
cavity perpendicular but edge slightly more rounded.
Aedeagus slightly longer than first three ventrites measured medially;
straight for apical three-fifths, thence gently curved to base, approximately
as wide as the antennal club excluding hairs.
Type material. - Paralectotypes, hereby designated, 3 females, labelled
“Key Largo, Fla.’’, ‘‘Beyer’’, ‘‘A. Fenyes Collection’’, and a label on one
pin in Fall’s hand ‘‘subtropicus Fall. Cotypes’’ (CAS).
There has never been a lectotype designated in spite of the ‘‘type’’ label
on the MCZ specimen. At the time of the Pandeleteius revision (Howden, 1959,
p. 388) a lectotype was unnecessary for nomenclatorial stability, but with the
description of a closely-related species, S. caymani, it is now essential.
. The lectotype is missing the right fore femur and tarsus and the left hind
tarsus and appears to be slightly teneral. It is 3.2 mm. long and1.3 mm. wide.
Distribution. - Cuba, southern Florida, Baja California. Total number of
specimens examined: 11. West Indian records: CUBA. 1 female, San Vin-
cente de Vinales, July, Archer (MCZ).
Discussion. - The specimen from Cuba is a female with the elytral swellings
much more pronounced than in more northern specimens.
The interrupted elytral swellings, slightly irregular striae, and form of the
aedeagus Should distinguish this species from all other West Indian tanymecines.
Even though males may have the elytral swellings greatly reduced, the striae
are disproportionately irregular in the vicinity of the swellings, making the
combination of these two characters useful for either sex.
In 1959 (pp. 388-389), I stated that subtropicus was incongruous with
Pandeleteius, but I did not recognize then that the more significant character
was in the modified ventrites rather than the short, slender aedeagus. In
Pandeleteius the anterior portion of ventrites 3, 4 and 5 is flat and unmodified.
7. Scalaventer caymani, new species
Figures 15, 40.
Diagnosis (based on females). - Elytral intervals 3, 5 and 7 slightly elevated
on disc, becoming flat before declivity. Sutural interval moderately swollen at
summit of declivity which is rather abrupt. Elytra parallel-sided.
Description. - Holotype, female, length 3.7 mm., width 1.4 mm. ; probably
somewhat teneral. Color pale cupreous and opalescent white with some dark
brown. Head mottled cupreous and white; pronotum cupreous with a broad white
vitta either side of median line; disc of elytra somewhat mottled, with a broad
dark brown fascia between about middle and apical third narrowly bordered
posteriorly by white. Ventrum and sides of beetle white. Swelling on sutural
interval at summit of declivity white, accenting it. Scales of irregular shapes,
contiguous (especially white scales) or not, strongly margined (except white
Howden: West Indian Tanymechini 23
scales), coarsely to finely granular, scales of head more strongly sculptured.
Setae of head and pronotum inconspicuous, mostly decumbent; setae of elytra
more conspicuous, longer than the average scale, completely or incompletely
arched, mostly uniserial on elytral intervals but more numerous on alternate |
intervals and sometimes multiserial where these intervals are most elevated.
In profile (Fig. 40), beak rather thick, its dorsum feebly arcuate; head
subspherical, very slightly more prominent between eyes. Head viewed an-
teriorly very robust, eyes scarcely exceeding it. Beak short, dorsal surface
one-tenth narrower than head between eyes, sides slightly divergent over
scrobes. Median line marked by a finely impressed line from about middle
of eyes to interantennal line and from thence obsoletely carinate. Interanten-
nal line glabrous for about half its length. Scales apicad of interantennal line
becoming sparser, smaller, smooth and convex; setae apicad of interantennal
line erect, becoming longer. Apical emargination of beak moderately shallow,
arcuate with its apex obtusely angled, occupying approximately one-half the
width of the beak apically; marked by an obsolete carina and with a row of
seven vibrissae on each side. Scrobe slightly obtusely angled, much wider at
angle, the vertical portion separated from apex by one scale and from eye by
four scales, slightly curved posteriorly, rapidly tapering and ending before
ventral surface below anterior edge of eye. Antenna with first segment of
funicle elongate, slender; segments 2 to 7 subequal, slender, about one-half
the length of first segment; antennal club moderate. Eye moderate, larger
than in subtropicus.
Prothorax broad, as wide as long, flattened, distinctly wider apically than
basally, sides feebly rounded between weak apical and basal constrictions.
Pronotum 1.8 times longer than prosternum, in profile 1.3 times longer dor-
sally than thick. Surface of pronotum with obsolete punctures between the
scales.
Elytra 2.6 times longer than prothorax, elytra across humeri 1.3 times
wider than prothorax. Elytra straight across base, humeri prominent, rather
abruptly right-angled, sides appearing parallel, but actually very slightly
divergent from base to apical three-fifths, thence gradually convergent to
apex, the slightly produced apical terminus of interval 5 interrupting the out-
line. Elytra in profile (Fig. 40) flat on basal fifth, thence very slightly arcu-
ate to declivity which is abrupt, slightly oblique. Sutural interval moderately
swollen at summit of declivity. Intervals 3, 5 and 7 slightly elevated from base
to approximately apical third, the elevations accentuated by more numerous
setae and more metallic color. Striae distinct, set with moderate, perfectly
aligned punctures.
Fore and hind legs subequal, middle legs definitely smaller. Fore femur
scarcely swollen, slightly bowed; fore tibia straight, slender, with five small
teeth on inner edge.
Fore coxae separated by distance equal to width of antennal club. Anterior
portion of ventrites 3, 4 and 5 (Fig. 15) abruptly, perpendicularly concave, the
modification slightly arcuate and widest at middle, the edge subcarinate.
Posterior-lateral angles of ventrites 2 and 3 slightly elongated, hooking over
the edge of the elytra. Ventrite 5 feebly convex along median line with a shallow
concavity on either side of median line at base.
Type series. - Holotype, female, Grand Cayman, W. Indies, 25 August
1908, Dr. M. Cameron, BM 1936-555, M. Cameron Journal W. I. 1178 (BM).
Paratypes, 2 females. GRAND CAYMAN: 1 female, same data as holotype
(BM); 1 female, June 1962, T. H. Farr (Inst. Jam.).
Discussion. - The two paratypes range in length from 3.5 to 3.6 mm, and
24 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
in width from 1.3 to 1.4 mm. Neither is as well marked as the holotype; in
one the elytral fascia is obsolete; in the other it is well-developed but less con-
spicuous because of additional dark brown spots over the disc. The extreme
plumpness of the beak in the type series could partly be attributable to particu-
larly robust specimens. There is no trace of the oblique ridges on the beak as
found in subtropicus. The prothorax is a little more convex in both paratypes
and the apical constriction is quite distinct on the disc in one. In both paratypes
the prothorax is very slightly longer than wide, thus lessening the relative width
compared to that of the base of the elytra; the length of the pronotum versus the
elytra is nearly the same as in the type.
In addition to the characters listed in the diagnosis, caymani also differs
from subtropicus in its flatter elytra with a dark fascia where subtropicus has
a white fascia; less swollen fore femora; larger eyes; and straight, perfectly
aligned strial punctures.
8. Scalaventer jamaicensis, new species
Figures 19, 66.
Diagnosis. - Elytral intervals equal, evenly setate. Elytra broadly rounded
apically. Aedeagus as long as ventrites 1 and 2 combined.
Description. - Holotype, male, length 2.5 mm., width1.0 mm. Specimen
with some scales abraded, missing tarsal claw on left middle leg, and aedeagus
cracked at tip. Color dark brown, tan and white; mottled, the only pattern
discernible being a vague pale ‘‘V’’ on apical third of elytra, this bordered an-
teriorly with dark. Scales contiguous or not, of irregular angular shapes,
strongly margined and strongly reticulate. Setae inconspicuous, small (about
as long as a scale), prostrate or slightly arched; uniserial on elytral intervals.
In profile, beak and frons flat to behind eyes, thence rather abruptly, obli-
quely angled to pronotum. Dorsally beak with sides slightly emarginate at base
where it is 0.81 times as wide as head between eyes, sides thence strongly
diverging apically, at apex as wide as head between eyes. Interantennal line
unmarked. Beak with a shallow basal concavity; median line finely carinate
from center of concavity to apex of epistoma, unmarked caudad of concavity.
Apex of beak slightly deflected; scales apicad of concavity gradually becoming
sparser, smaller, smoother and more convex; setae apicad of concavity erect,
becoming longer. Apical emargination of beak arcuate, its apex obtusely
angled; emargination occupying two-thirds of beak apically, marked by a weak
carina and with two slender vibrissae on each side of base.
Epistoma with median line finely carinate. Scrobe obtusely angled, wider at
angle; the vertical portion at its middle about equidistant between eye and apex,
slightly tapered, curved, ending just before ventral surface beneath anterior
edge of eye. Antenna with scape slightly bowed; club moderate, compact. Eye
moderate, quite convex, protruding well beyond sides of head.
Prothorax as long as wide, sides gently rounded between basal and apical
constrictions. Pronotum 1.7 times longer than prosternum; in profile 1.25
times longer dorsally than thick. Surface of pronotum with obsolete punctures
between the scales.
Elytra 2.3 times longer than prothorax, elytra across humeri 1.2 times
wider than prothorax. Base of elytra slightly arcuately emarginate. Elytra
rather strongly convex transversely. Elytra with sides slightly divergent from
humeri to middle, thence broadly rounded to apex; the broadly rounded apical
terminus of intervals 4 to 6 interrupting the outline. Elytra in profile (Fig. 19)
Howden: West Indian Tanymechini 25
distinctly arcuate from base to apex, highest medially; declivity slightly indi-
cated, oblique, slightly concave. Elytral intervals equal, slightly convex
especially basally and medially. Striae set with moderate, perfectly aligned
punctures.
Fore leg only slightly longer than other legs; fore femur scarcely swollen;
inner edge of fore tibia with only one distinct denticle and some serrulation.
Fore coxae separated by distance equal to approximately three-fifths the
width of the antennal club. Anterior portion of ventrites 3, 4 and 5 abruptly,
perpendicularly concave, the modification slightly wider medially, the edge
not carinate. Ventrite 5 slightly convex, its apex broadly rounded and witha
small median emargination.
Aedeagus (Fig. 66) as long as ventrites 1 and 2 combined, slightly arcuate,
approximately four-fifths as wide as antennal club; apical opening elongate-
elliptical, its apex briefly truncate.
Allotype, female, length 3.2 mm., width 1.4 mm. Specimen with left fore
tibia and tarsus mounted on point. Color and pattern similar to holotype.
Differs from holotype in the following respects. Beak in profile thicker; head
and beak more robust. Beak with carina of median line obsolete. Apex of
beak less deflected than in holotype. Apical emargination deep, arcuate, its
apical angle obsolete; epistoma with median line obsoletely carinate; a row of
four vibrissae on each side of apical emargination, and two small vibrissae
arising from epistoma itself. Elytra 2.5 times longer than prothorax. Elytra
with sides broadly rounded from about basal sixth to apical terminus of inter-
vals 4 to 6, the apex slightly attenuated beyond this. Elytra in profile with
declivity more evident, its summit well below the highest part of the elytra due
to their convexity. Sutural interval somewhat enlarged at summit of declivity.
Fore tibia with four denticles on inner edge. Fore coxae separated by distance
equal to width of antennal club. Anterior modifications of ventrites as in holo-
type but more abrupt. Ventrite 5 nearly flat, its apex narrowly rounded.
Type series. - Holotype, male, Port Henderson, St. Catherine Parish,
Jamaica, 28 September 1945, E. L. Sleeper Collector (Long Beach). Allo-
type, female, Port Henderson Hill, St. Catherine Parish, Jamaica, 11 March
1959, T. H. Farr (Inst. Jam.). Paratypes, 2 males, 4females. JAMAICA:
1 female, same data as holotype (Long Beach); 1 male, 2 females, same data
as allotype (Howden, Inst. Jam.); 1 male, Ewarton, St. Catherine, 28 Sep-
tember 1945, E. L. Sleeper Collector (Sleeper); 1 female, Portland Ridge,
Clarendon, 6 July 1962, T. H. Farr (Inst. Jam.).
Discussion. - Males vary in length from 2.3 to 2.5 mm. and are 0.9 mm.
wide; females vary in length from 2.3 to 3.2 mm. and in width from 1.1 to 1.4
mm. The only additional color pattern evident in the paratypes is a dark medi-
an vitta on the vertex. One female paratype has the scales neither margined
nor reticulate but granular. The beak is usually as described for the holotype
rather than the allotype. Average number of denticles and teeth on the inner
edge of the fore tibia is three. The elytra are variable in relative width
giving them a disconcertingly different habitus; at their widest point the elytra
vary from 1.1 to 1.4 times wider than their humeri. The declivity of the
females in profile also is variable; in two paratypes it is as in the allotype and
in the other two is less well-marked. The anterior modification of ventrite 5
is subcarinate in one female. In the males the apex of ventrite 5 is not
emarginate in one specimen and obsoletely emarginate in the other.
Scalaventer jamaicensis seems most closely related to caymani (known
from Grand Cayman) and convexifrons (known from the Bahama Islands),
though geographically it is more apt to be confused with coccolobae. From
26 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
convexifrons, jamaicensis may be separated by its much shorter aedeagus and
shorter, broader elytra. From caymani (and subtropicus) it is readily dis-
tinguished by its nearly flat, even elytral intervals. From coccolobae, jamai-
censis may be separated by its more widely separated fore coxae, scarcely
enlarged fore femora and noncarinate ventral modifications. In habitat, jam-
aicensis is closest to coccolobae; jamaicensis is known from low hills in south-
ern Jamaica and coccolobae from low hills over much of the island, the known
ranges being most approximate west of Kingston.
9. Scalaventer convexifrons, new species
Figures 21, 58, 67.
Diagnosis (based on male). - Aedeagus as long as entire abdomen. In pro-
file, head prominently swollen above eyes.
Description. - Holotype, male, length 3.0 mm., width 1.1 mm. Color
black, white and tan, marked as follows: sides of beak and head white except
for broad dark vitta behind eye to pronotum, remainder of head mottled; pro-
notum with a white vitta either side of median line; elytra with intervals 2 and
3 white on basal fifth; elytra with an irregular, conspicuous, white ‘‘V’’ on
apical half, bordered anteriorly with black; intervals 5, 7 and 8 black on de-
clivity before apex; remainder of elytra including epipleurae mottled; legs
indistinctly annulate. Scales angular, contiguous or not, obsoletely to finely
margined, granular; white scales less strongly sculptured but often subpustu-
late towards declivity. Setae of frons and prothorax completely, slightly arched,
as long as one to one and a half scales; setae of head behind convexity prostrate,
very inconspicuous; setae of elytra becoming incompletely arched towards side
and apex; uniserial on elytral intervals.
In profile (Fig. 58), beak and frons obsoletely concave to above middle of
eye where head is prominently convex, thence flat, oblique (slightly more than
45 es slightly elongate to pronotum. Beak with sides Slightly converging from
over insertion of antennae to base where it is 0.92 times as wide as head be-
tween eyes; sides rapidly, briefly converging apicad of insertion of antennae,
thence parallel for a short distance. Dorsum of beak obsoletely concave, me-
dian line finely impressed from base to between middle of eyes. Interantennal
line unmarked, apex of apical emargination reaching interantennal line. Apical
emargination right-angled, occupying approximately three-fifths of the apical
edge of beak; emargination obsoletely carinate, except at apex where it is
slightly elevated, a fine carina extending from apex to base; with a row of six
relatively short vibrissae each side of emargination; apical edge of beak re-
flexed and broadly glabrous either side of emargination. Scrobe obtusely
angled, wider at angle, the vertical portion about equidistant between eye and
apex, tapering rapidly and ending well above ventral surface beneath anterior
edge of eye. Antenna with scape slightly bowed; club moderately large, elon-
gate, slightly ‘‘loose’’ because of slight constriction at base of segments 2 and
3. Eye moderately small, quite convex, well separated from dorsal surface,
approximately equidistant between dorsal surface, ventral surface and pro-
thorax.
Prothorax as long as wide, sides rounded between constrictions, apical
constriction 1.2 times wider than basal constriction. Pronotum 1.6 times
longer than prosternum; in profile, 1.2 times longer dorsally than thick.
Punctures obsolete on disc becoming moderate on sides.
Elytra 2.4 times longer than prothorax, elytra across humeri 1.3 times
Howden: West Indian Tanymechini 27
wider than prothorax. Elytra nearly straight across base. Elytra with sides
parallel to middle, thence gradually converging to apex, the apical terminus
of intervals 4 to 6 scarcely interrupting outline. In profile (Fig. 21), disc of
elytra weakly arcuate from base nearly to apex, the summit of declivity obso-
lete, the declivity oblique, straight. Elytral intervals equal, very slightly con-
vex beyond basal third. Striae set with moderately small punctures, slightly
irregularly aligned on basal third.
Fore legs only slightly longer than other legs; fore femur scarcely more
swollen than hind femur; fore tibia straight, inner edge with four distinct,
moderate teeth on right tibia, five on left tibia.
Fore coxae separated by distance equal to the greatest width of the anten-
nal scape, approximately one-half the width of the antennal club. Anterior
modification of ventrites 3, 4 and 5 abrupt, perpendicular, slightly wider
medially; gradually becoming more elevated medially, the edge not carinate.
Ventrite 5 rather strongly convex longitudinally, apex rounded.
Aedeagus (Fig. 67) as long as entire abdomen measured medially, about as
thick as thickest part of scape, slightly more strongly arcuate for basal half;
apical opening short, oval, apex briefly truncate.
Type series. - Holotype, male, West side Port Nelson, Rum Cay, Bahama
Islands, 6 April 1965, B. D. Valentine, R. W. Hamilton Collectors (AMNH,
temporarily Valentine). No paratypes.
Discussion.- Scalaventer convexifrons appears to be most closely related
to jamaicensis which is particularly similar in its head and beak. However,
convexifrons has the head much more swollen than any other Scalaventer and
consequently the eye approximately equidistant between the dorsal and ventral
surfaces. Note that the head and prothorax are slightly tilted to the right side
in the photograph (Fig. 21), slightly distorting the true profile; figure 58 de-
picts the profile accurately. The much longer aedeagus of convexifrons is, of
course, a more decisive character, but for females the swelling over the eyes
and narrower, longer elytra will probably readily distinguish convexifrons
from jamaicensis. Series of convexifrons are needed to determine whether or
not the larger and more numerous tibial teeth and the shorter, narrower scrobe
are reliable characters.
The swollen frons (Fig. 58) and therefore the eye remote from dorsal sur-
face are generic characters of Pandeleteinus (Fig. 57); in this context the criti-
cal characters of S. convexifrons are: scrobe not reaching ventral surface,
eye distant from pronotum, anterior modification of ventrites 3, 4 and 5 strong-
ly developed, and fore coxae relatively widely separated.
Geographically, convexifrons is the most remote of all the Scalaventer and
is the only species known from the Bahama Islands.
10. Scalaventer gelinasus, new species
Figures 9, 11, 22.
Diagnosis (based on female). - Beak very short and robust; apex abruptly
deflected so that epistoma is perpendicular to dorsum; scrobes encroaching
upon dorsum so that at narrowest point dorsum is only half as wide as head be-
tween eyes.
Description. - Holotype, female, length 3.7 mm., width 1.4 mm. Speci-
men teneral with mandibular cusps still attached. Color testaceous because of
teneral condition, only white markings distinct. Marked with white in a front-
al spot, four small spots across middle of prothorax, a faint ‘‘V’’ on apical
third of elytra and a spot on sutural interval at summit of declivity. Scales
28 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
finely to obsoletely margined, coarsely to finely granular. Setae as long as
one and one-half scales; completely, slightly arched, becoming incompletely
arched towards apex and sides; more numerous on elytral intervals 3, 5 and 7.
In profile, head and beak very short and robust, apex of beak abruptly
truncated; beak thicker than long; dorsum of head and beak flat, gently arcuate
over caudal half of eyes. Dorsal surface of beak (Fig. 9) apically reduced in
size, widest over insertion of antennae where sides of beak are 1.5 times
wider than dorsum; narrowest at base where the converging apices of the
scrobes reduce dorsum to 0.5 times as wide as head between eyes and where
sides of beak are 2.4 times wider than dorsum; apical edges of dorsum strong-
ly converging from insertion of antennae to apex of epistoma, the sides here
falling off rapidly; median line deeply impressed from interantennal line to be-
yond middle of eyes. Interantennal line unmarked. Apical emargination ap-
proximately right-angled, carinate, the carina extending caudally for a short
distance; with a row of seven conspicuous, apically-directed vibrissae on each
side; epistoma small, nearly perpendicular to dorsum, entire anterior edge of
epistoma slightly arcuately emarginate. Scrobe with horizontal portion almost
non-existent; vertical portion strongly bowed towards apex of beak, slightly
closer to apex of beak than to eye, ending before ventral surface in front of
anterior edge of eye. Scape strongly bowed to accommodate curvature of beak.
Antennal club moderate in size, first segment with slightly elongate base. Eye
moderately small, moderately convex.
Prothorax 1.1 times longer than wide, sides gently rounded between con-
strictions; basal constriction close to base. Pronotum 1.8 times longer than
prosternum; in profile disc slightly arcuate between constrictions, apical con-
striction obsolete on disc. Punctures small, moderately deep.
Elytra 2.6 times longer than prothorax; elytra across humeri 1.4 times
wider than prothorax. Base of elytra nearly straight. Elytra with sides
parallel for basal seventh, thence slightly divergent to beyond middle, thence
convergent, gently rounded to apex, apical terminus of intervals 4 to 7 moder-
ately conspicuously interrupting outline; apex broadly, slightly truncate. In
profile (Fig. 11), elytra very slightly, evenly arcuate to summit of declivity
which is rather broadly rounded; declivity slightly oblique. Basal seventh of
intervals 2 to 5 conjointly convex. Intervals 3, 5 and 7 slightly more elevated
and wider apicad of basal seventh, otherwise intervals equal on disc. Striae
set with small, moderately deep punctures.
Fore leg slightly larger than hind leg; fore femur with swelling moderately
weak, gradual. Inner edge of fore tibia with six moderate to large teeth and a
few minute denticles.
Fore coxae separated by distance approximately equal to the width of the
antennal club. Abdomen (Fig. 22) with modification of anterior portion of
ventrites 3, 4 and 5 deep, but oblique rather than perpendicular to surface,
edges not carinate, slightly arcuate for central two-thirds in ventrites 3 and 4,
less arcuate in ventrite 5. Ventrite 5 strongly, longitudinally convex for its
entire length; apex deflected, rather narrowly rounded.
Type series. - Holotype, female, Loma Rucilla and mountains North,
Dominican Republic, June 1938, 5-8000 feet, Darlington (MCZ). No paratypes.
Discussion. - Variation may be expected (particularly in males) in color
and color pattern, relative robustness of the head and beak, and contour of
declivity. Females will probably show some variation in the abdominal modi-
fications, but the modifications will probably always be slightly arcuate, obli-
que and not carinate. The longitudinal convexity of ventrite 5 of females is
undoubtedly stable, and this character is unique for the genus.
Howden: West Indian Tanymechini 29
S. gelinasus may be distinguished from other Scalaventer by the following
characters: extremely short, robust beak and consequently with horizontal
portion of scrobes nearly absent, dorsal surface strongly constricted basally
to one-half the width of head between eyes, scape strongly bowed, epistoma
perpendicular to dorsum of beak; slightly oblique declivity; females with
modification of anterior portion of ventrites 3, 4 and 5 slightly arcuate and not
carinate, ventrite 5 longitudinally convex, the convexity extending to sides.
Of other West Indian tanymecines with a dorsally strongly constricted beak,
Isodrusus has connate claws and S. valkyrius has the summit of the elytral
declivity produced, elytral intervals 2 and 4 without setae on disc, and the
female abdominal modification carinate.
S. gelinasus is the only species of Scalaventer known from Hispaniola.
Other tanymecines on the island are Polydacrys with semi-enclosed corbels
and stalked mandibular scars and three Paululusus species which have scarcely
modified ventrites. S. gelinasus has the weakest modification of the ventrites
in the genus, but, as is seen in the photograph (Fig. 22), the modification ex-
tends the entire width of the abdomen and when compared to Paululusus is
strong indeed.
11. Scalaventer valkyrius, new species
Figures 12, 23, 24.
Diagnosis (based on female). - Beak very short, horizontal portion of
scrobe greatly reduced. Sutural interval produced posteriorly at summit of
declivity.
Description. - Holotype, female, length 2.6 mm., width 1.0 mm. Color
black, white and cupreous, marked as follows. Head cupreous with dark
triangle on vertex. Pronotum predominantly cupreous, faintly mottled. Ely-
tra bright cupreous; sutural interval darker, somewhat mottled, the dark area
extending to stria 4 on basal seventh; a conspicuous white lunule on intervals
5, 6 and 7 at basal third, the lunule broadly bordered in black; a slightly obli-
que, broad, white fascia on apical third bordered anteriorly with black. Scales
of various shapes, often rounded, not contiguous, strongly margined, coarsely
granular, Setae of head and pronotum small and inconspicuous except between
eyes and on apex of pronotum where they are completely arched, scarcely
elevated, as long as one and one-half to two scales. Setae of elytra completely
arched but higher than those of head and pronotum, as long as two scales;
setae of summit of declivity incompletely arched; setae uniserial; absent on
intervals 2 and 4 except on declivity, sparse on interval 6.
In profile, beak and head flat from apex to over eye where it is gradually,
gently rounded, thence less rounded to pronotum; beak thicker than long (Figs.
12, 23). Beak very short, approximately half as long as distance between
eyes. Dorsum of beak obsoletely concave, median line briefly marked with
Slight groove; dorsal edge of beak excavate over scrobe, at its narrowest point
0.7 times narrower than head between eyes. Interantennal line unmarked; api-
cad of line beak deflected, without scales, with straight, slender vibrissae.
Apical emargination slightly obtusely angled, occupying approximately one-
half the apical edge of the beak, finely carinate, a short, fine carina extending
from apex to interantennal line; with a row of three vibrissae on each side of
emargination. Scrobe broad at angle, antenna inserted above anterior edge of
vertical portion, the horizontal portion extending anteriorly only very briefly
beyond insertion; dorsal edge of horizontal portion strongly sinuate; vertical
30 Contrib. Amer. Ent. Inst., vol. 55 noid, 1970
portion slightly bowed, about equidistant between eye and apex at its narrow-
est point, deep and well-defined to its termination on ventral surface beneath
anterior edge of eye. Scape slightly bowed to accommodate curvature of beak.
Antennal club moderate in size, first segment with slightly elongated base. Eye
moderately small, only moderately convex, in anterior view only slightly ex-
ceeding the robust head.
Prothorax 1.1 times longer than wide, sides gently rounded between con-
strictions. Pronotum 2.0 times longer than prosternum; in profile, disc
slightly arcuate between constrictions, the apical projection nearly three times
as long as the basal. Punctures small, shallow on disc, becoming moderate on
Sides.
Elytra 2.3 times longer than prothorax, elytra across humeri 1.3 times
wider than prothorax. Base of elytra slightly, arcuately emarginate between
striae 5. Elytra with sides weakly divergent to about middle, thence con-
vergent, gently rounded to apex, the apical terminus of intervals 4 to 6 obso-
letely interrupting outline. In profile (Fig. 23), elytra nearly flat for basal
fifth, thence very slightly arcuate to apex of declivity; declivity with approxi-
mately dorsal third directed slightly, obliquely inwards, remainder directed
slightly apically, the apex of elytra scarcely exceeding the apex of the declivity.
In posterior view elytra rather strongly convex. Elytral striae set with small
punctures; elytral intervals nearly equal, intervals 3 and 5 obsoletely enlarged
on disc.
Fore leg slightly larger than hind leg; fore femur moderately swollen, the
swelling gradual basally, moderately abrupt distally. Fore tibia slender
throughout, slightly bowed, with six minute denticles on inner edge.
Fore coxae separated by approximately the width of the antennal club. Ab-
domen with posterior-lateral edge of ventrite 2 slightly enlarged and overlap-
ping edge of elytra. Modification of anterior portion of ventrites 3, 4 and 5
(Fig. 24) deep throughout, but arcuate for only approximately median two-thirds,
slightly elevated and carinate, the modification strongest on ventrite 5. Ven-
trite 5 flattened, slightly convex basally, apex broadly rounded.
Type series. - Holotype, female, Hardwar Gap, Jamaica, 4000 feet, 29
July 1966, A. T. Howden, not on Cyrilla (Howden). No paratypes.
Discussion. - When additional specimens of this species are found, they
will probably exhibit variation in the color pattern, general contours of the
elytra and details of the vestiture. Males may have a more slender head,
slightly larger eye, more enlarged fore leg, more closely placed fore coxae,
and possibly a reduced swelling at the summit of the declivity.
S. valkyrius can be distinguished by the following combination of characters:
beak very short, horizontal portion of scrobe greatly reduced; sutural interval
produced posteriorly at summit of declivity; intervals 2 and 4 on disc without
setae; ventrite 5 of female with anterior modification strongly arcuate and
elevated on central two-thirds. S. remotus has a similar declivity and strik-
ingly similar color pattern, but has all intervals setate and a long beak with
long, gently angled scrobe. S. litoreus and montanus females have the modifi-
cation of ventrite 5 arcuate but they are never as strongly arcuate or as strong-
ly carinate as in valkyrius; litoreus and montanus may also be distinguished
from valkyrius by their robust form and broad, flat beak with gently curved
scrobe. The abdomen of Isodrusus guajavus is very close to that of S. valkyri-
us, and both species also have similar beaks and scrobes; the connate claws
and narrower beak of Isodrusus will immediately separate it from Scalaventer.
The holotype was taken at the end of a month’s collecting at Hardwar Gap
when a particular effort was made to collect weevils on trees other than the
Howden: West Indian Tanymechini 31
common hosts of Scalaventer.
12. Scalaventer cubensis, new species
Figures 18, 41, 42, 64.
Diagnosis (based on male). - Dorsum of beak long, narrow, parallel-sided,
gradually deflected apicad of interantennal line, scales typical nearly to apex.
Apical emargination small, occupying only one-third of apical width of beak.
Description. - Holotype, male, length 3.1 mm., width1.2 mm. Color
pale tan; elytra slightly mottled, sutural interval somewhat cupreous from
basal fifth to apex. Scales of irregular shapes, mostly angular, contiguous,
finely to moderately granular, obsoletely to moderately margined; scales of
sutural interval becoming pustulate about middle, some additional pustulate
scales on declivity especially on alternate intervals. Setae completely arched,
scarcely elevated, as long as one and one-half to two scales; setae of elytra
uniserial and evenly distributed except on sutural interval at summit of declivi-
ty where they are more numerous.
In profile (Fig. 41), beak gently deflected apicad of insertion of antennae,
flat to above middle of eyes, thence gently, evenly arcuate to pronotum. Head
(Fig. 42) narrow between eyes; beak approximately same width as head be-
tween eyes, relatively long and narrow, sides nearly parallel. Beak 1.2 times
wider than long. Median line rather deeply impressed from interantennal line
to middle of eyes; dorsum of beak slightly longitudinally convex either side of
midline. Interantennal line unmarked. Scales apicad of interantennal line
gradually becoming smooth and convex but scarcely less dense than typical
scales. Apical emargination small, approximately right-angled, occupying
approximately one-third of apical edge of beak, apex well removed from inter-
antennal line; emargination marked by a very weak carina and a single vibrissa
on either side. Scrobe with dorsal and posterior edge forming a right-angle;
horizontal portion almost as long as vertical portion; very wide at angle; verti-
cal portion tapering rapidly to a point closer to eye than to apex of beak, ending
well above ventral surface beneath anterior edge of eye. Scape gradually
thickened from base, very slightly bowed; antennal club moderate, compact.
Eye moderately large, moderately convex, facets relatively small.
Prothorax 1.2 times longer than wide, sides only slightly rounded between
constrictions. Pronotum 1.6 times longer than prosternum; in
profile, 1.5 times longer dorsally than thick, disc slightly arcuate. Surface
with punctures shallow on disc becoming moderately deep on sides. Anterior
edge of pronotum with a compact row of seven anteriorly-directed scales at
the site where ocular vibrissae are situated in continental tanymecines.
Elytra 2.1 times longer than prothorax, elytra across humeri 1.3 times
wider than prothorax. Elytra approximately straight across base. Elytra
with sides parallel for basal fifth, from there very slightly diverging to just
beyond middle, thence rounded to apex, the apical terminus of intervals 4 to
6 broadly interrupting outline. Elytra in profile scarcely arcuate; summit of
declivity broadly rounded, declivity slightly oblique, slightly concave. Inter-
vals 3, 5 and 7 obsoletely elevated, otherwise intervals even. Striae closely
set with moderately deep, perfectly aligned punctures.
Legs long and slender, fore legs longer but relatively scarcely thicker.
Fore tibia with six distinct, large teeth on inner edge.
Fore coxae separated by distance equal to approximately four-fifths the
width of the antennal club. Abdomen densely squamose, the scales little differ-
32 Contrib, Amer. Ent. Inst. , vol. 5,0. 5, 1970
ent from those of dorsum. Anterior portion of ventrites 3, 4 and 5 (Fig. 18)
perpendicularly concave, deep throughout, the edge not carinate, the squamose
portions slightly elevated anteriorly medially. Ventrite 5 moderately convex,
its apex truncate-emarginate.
Aedeagus (Fig. 64) gently, evenly arcuate, as long as first two and one-
half ventrites measured medially, thickened apically; apical opening broadly
elliptical. |
Type series. - Holotype, male, Coast below Pico Torquino, Cuba, 26-30
June 1936, Darlington (MCZ). No paratypes.
Discussion. - In females the beak will probably be wider, on the sides
especially, but the rounded apex with its nearly typical scales and very small
epistoma should be fairly constant. The moderately deep strial punctures of
the type may be considerably moderated in females. It will be interesting to
see the condition of the ocular vibrissae site in additional specimens.
S. cubensis appears to most closely related to jamaicensis, by virtue of its
aedeagus being shorter than the combined length of ventrites 1 to 3; its scarce-
ly enlarged fore legs; its short, truncated elytra; and its nearly equal elytral
intervals. From jamaicensis, cubensis may be separated by its much smaller
epistoma, narrower beak, more strongly dentate fore tibiae, and shorter api-
cal opening of the aedeagus.
Paululusus, new genus
Size small, usually between 2 and 3 mm. in length. Body and legs covered
with scales and small setae. Beak deflected from head by less than 90°, slight-
ly wider than long, dorsal surface flattened, sometimes slightly concave thence
slightly convex apicad of interantennal line which may or may not be marked by
a short glabrous line. Apex of beak triangularly emarginate, the emargination
marked by a fine carina posteriorly and occupying between one-third and two-
thirds of the width of the beak anteriorly; apical edge of epistoma emarginate.
Sides of beak nearly vertical; scrobes deep, obtusely or nearly right-angled,
not reaching ventral surface, the vertical portion approximately equidistant
between eye and apex of beak. Antennal scape without scales, reaching middle
of eye; funicle seven-segmented; first segment longer than others which are
subequal or equal; club short to moderate, narrowly to broadly elliptical.
Frons not swollen in lateral profile. Eye small to moderate in size, strongly
convex and protruding, nearly hemispherical in some males, situated at extreme
anterior edge of head close to dorsal surface. Prothorax a little longer than
wide, feebly convex between basal and apical constrictions which are moderate
on the sides, weaker on the disc. Pronotum produced anteriorly over head; api-
cal margin without ocular vibrissae, teeth or lobes. Elytra with humeral ang-
les well-developed or not. Stria 10 obsolete medially. Fore leg larger than
other legs; fore femur swollen or not, fore femur without teeth or serrations
on inner edge; fore tibia dentate on inner edge. Fore coxae separated by a
distance equal to approximately one-third to one-half the width of the antennal
club, more widely separated in female than in male. Hind tibia with corbel
open. Tarsi completely padded; tarsal claws free. Abdomen with the anterior
portion of ventrites 3, 4 and 5 weakly modified, at most narrowly glabrous and
narrowly concave. Aedeagus very slender and longer than the first four ven-
trites measured medially, sometimes longer than all five ventrites.
Type-species. - Paululusus calypso new species.
The name Paululusus is masculine and refers to the miniature appearance
Howden: West Indian Tanymechini 33
of the insects.
Discussion. - Paululusus is most closely related to the continental neo-
tropical genus Pandeleteinus Champion. The major differences between the
two genera are as follows. Paululusus (Fig. 51) has the frons not swollen, the
head evenly arcuate in profile, eyes close to dorsal surface, ocular vibrissae
absent, scrobe not reaching ventral surface, and fore coxae separated by one-
third to two-thirds the width of the antennal club. Pandeleteinus (Fig. 57) has
the frons transversely swollen, the head and frons consequently usually sinuate
in profile; eyes distant from dorsal surface; ocular vibrissae present; scrobe
reaching ventral surface (except in submetallicus (Schffr.)); fore coxae contigu-
ous or very narrowly separated.
In addition, there are several secondary distinguishing characteristics.
The dorsal surface of the tarsi, the articulating surface of the corbels and, to
a lesser extent, the scape are all densely squamose in Pandeleteinus; in Paulu-
lusus there are no scales or only scattered scales in these places. The size of
Pandeleteinus ranges from 2.7 to 4.5 mm; Paululusus ranges from 2.0 to 3.2
mm. The aedeagus of Pandeleteinus is often as long proportionately as in
Paululusus but it is generally thicker. Pandeleteinus always has a much broad-
er beak than Paululusus; this is too difficult to describe objectively to be use-
ful, but when specimens are available for comparison, the difference is con-
Spicuous. Pandeleteinus is known from southwestern United States and Mexico.
Paululusus is known only from Hispaniola.
Paululusus is also related to Isodacrys, the apterous Paululusus constanzae
n. sp. in particular resembling an Isodacrys. In Paululusus the apterous con-
dition is a specific and not a generic character. Isodacrys is known only from
the continental Neotropics and is characterized by short, fused elytra without
humeri and by the absence of wings. Isodacrys differs from Paululusus in: its
scrobe reaching the ventral surface; aedeagus much stouter; beak broader and
parallel-sided; tarsi, scape and corbels densely squamose; and ventrites more
strongly modified. For a discussion of these genera in reference to the apter-
ous condition see the Introduction.
Paululusus differs from Scalaventer in having the modification of the anteri-
or portion of ventrites 3, 4 and 5 never abruptly concave across the entire width
of the abdomen and never with the edge carinate.
Nothing is known of the ecological requirements of the species of Paululusus
other than that revealed by the labels on the specimens. Paululusus constanzae
was collected in the remote interior mountains between 3000 and 5000 feet;
calypso and hispaniolae are known from numerous localities at sea level up to
about 2000 feet, along the coast and in the interior (see Map 4). One species,
hispaniolae, was collected on ‘‘calmite’’. Specimens of calpyso and hispaniolae
were collected together by beating branches in a semi-dessert area; calypso was
also taken by sweeping a weedy, bushy hillside. One specimen of hispaniolae
was collected on ‘‘flowering plants’’. This indicates that adults of Paululusus
like those of Isodacrys and Pandeleteinus feed principally on herbaceous plants
and bushes, whereas Scalaventer and Pandeleteius feed principally on trees.
Key to the Species of Paululusus
1. Elytra with setae inconspicuous, completely arched; sutural interval swol-
len at sunimit ofdeclivity: declivity abrupt... ac ore a ee Pa es
Se ges as at ee. eR et ee geen 2. hispaniolae, new species (p. 36)
Elytra with setae conspicuous, erect; sutural interval not swollen at sum-
mit of dechivitvs Geclivity evadtar. sa es es ke ee ee ee 2
34 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
2. Humeri well-developed; elytra not fused; wings fully developed. ....
“i TM Ra aye » 2 ees. 1. calypso, new species (p. 34)
Humeri absent; elytra fused? wings absentin yy. eae
chee ae Ge ... 93 constanzae, new species (p. 38)
1. Paululusus calypso, new species
Figures 27, 29, 54, 68.
Diagnosis. - Elytra with well-developed humeri; elytral intervals each
with a row of erect setae.
Description. - Holotype, male, length 2.2 mm., width 0.7 mm. Body and
legs covered with pale brown and pale green scales; the dorsum predominantly
pale brown, the sides of head, thorax and elytra mostly green. Marked with
brown in a broad median thoracic vitta which continues, tapering over head to
between eyes; elytra with a short basal vitta on intervals 5 and 6. Scales of
dorsum and legs not contiguous, finely granular, not margined, of irregular
polygonal shapes; scales of head, beak and tibiae more rounded in shape.
Scales of declivity with a metallic lustre. Scales of ventrum pale green or
opalescent, contiguous or overlapping, rounded in shape. Elytral intervals
each set with a row of short, blunt, erect setae; setae of elytral declivity much
more numerous; setae of beak like those of elytra, numerous; setae of frons
semi-erect; setae of head and pronotum inconspicuous, fine and nearly pro-
strate.
In profile, head sub-spherical, beak rather thin, feebly arcuate over scrobes.
Beak slightly narrower than head between eyes, with sides approximately paral-
lel; dorsal-lateral edge rounded between head and antennal insertion, abrupt and
slightly convex over horizontal portion of scrobe. Beak obsoletely concave
from base to interantennal line, thence feebly convex; interantennal line marked
only by a brief glabrous area either side of median line. Median line of beak
marked with a feeble, glabrous carina from apex to interantennal line from
whence it is marked by a fine, deep groove terminating beyond middle of eyes,
almost contiguous with the dark median vitta of vertex. Apical emargination
approximately right-angled, occupying approximately one-half of the width of
beak apically and marked posteriorly by a fine but distinct carina. Scrobe
deep, obtusely angled, vertical portion closer to eyes than to apex of beak,
feebly curved posteriorly and ending beneath anterior edge of eye before the
ventral surface. Antenna with scape straight, clavate; funicle with first seg-
ment robust, conical; segments 2 to 7 subcylindrical, increasingly broader
and shorter; club elliptical with apex acute. Eye moderate in size, strongly
convex, thickest posteriorly.
Prothorax 1.05 times longer than wide, sides gently rounded between con-
strictions. In profile pronotum slightly convex; pronotum 1.6 times longer
than prosternum. Surface of pronotum between constrictions with small, deep
foveae separated from each other by one scale.
Elytra 2.4 times longer than prothorax; elytra across humeri 1.3 times
wider than prothorax. Elytra with humeri rounded, right-angled, base obso-
letely, arcuately emarginate; sides of elytra slightly divergent from humeri to
middle thence rounded to apical eighth, the apex attenuate beyond this. Elytra
in profile with disc rising slightly from base to beyond middle, thence gently
rounded to apex, the declivity weak, vague; epipleura slightly emarginate op-
posite hind coxa. Elytral striae not prominent, set with small punctures which
are separated from each other in the striae by one scale. Sutural interval
Howden: West Indian Tanymechini 35
narrow, as wide as one scale; other intervals equal, as wide as two to three
scales; all intervals flat or nearly so, except interval 10 on apex.
Fore leg definitely longer than hind leg, middle leg definitely shorter than
hind leg. All legs with femora swollen medially, tibiae slender, tarsi long.
Fore leg (Fig. 54) with femur grossly, abruptly swollen; tibia with four teeth
and several denticles on inner edge.
Fore coxae separated by a distance equal to approximately one-third the
width of the antennal club. Anterior portion of ventrites 3, 4 and 5 narrowly
glabrous across their entire width, slightly concave medially. Ventrites 2, 3
and 4 with their posterior margins slightly convex. Ventrite 5 slightly convex,
its apex without scales and set with many fine hairs, its apical margin trun-
cate and slightly emarginate medially.
Aedeagus (Fig. 68) bent in type, approximately as long as entire abdomen.
Aedeagus very slender, about as wide as thickest part of scape; cylindrical;
apical three-fifths straight, basal two-fifths gently arcuate; apex in lateral
view Slightly less than a 45° angle; apical opening in dorsal view elliptical.
Allotype, female, length 2.8 mm., width 1.1 mm. Similar to holotype
except in the following respects: head and beak (Fig. 29) more robust especial-
ly in profile; sculpture of beak slightly weaker. Prothorax approximately 1.2
times longer than wide. Elytra (Fig. 27) much more attenuate, 2.7 times
longer than pronotum. Legs scarcely different from holotype. Fore coxae
more widely separated, i.e., by approximately half the width of the antennal
club. Anterior portion of ventrites 3 and 4 glabrous across their entire width
but in only a thin line, the glabrous area scarcely concave. Ventrite 5 elongate
its sides converging from base and its apex narrowly rounded; convex along
median line and with a large depression on either side; only 12 atypical scales
on convexity; the surface granular and set with fine hairs as long as the elytral
setae.
Type series. - Holotype, male, Port au Prince, Haiti, 1 March 1908,
Dr. M. Cameron, B. M. 1936-555 (BM). Allotype, female, same data as
holotype (BM). Paratypes, 16 males, 19 females. HAITI: 5 males, 14 fe-
males, same data as holotype (BM, Howden); 1 female, Diquini, W. M. Mann
(MCZ); 1 female, 10 Km. N. of Ennery, 1000 feet, 12 July 1956, B. and B.
Valentine, sweeping on bushy, weedy hillside (Valentine); 1 female, Hayti,
Parish, 1899, Sharp Coll. 1905-313 (Howden); 10 males, 32 Km. SW Mirebalais,
c500 feet, 5 July 1956, B. and B. Valentine, beating branches in semi-desert
(Valentine, Howden); 1 male, Petionville, 1000 feet, 7 July 1956, B. and B.
Valentine, sweeping on bushy hillside, late morning (Valentine); 1 female,
Port au Prince, R. J. Crew, Wickham Coll. 1933 (USNM); 1 female, Moun-
tains near Port au Prince, up to 2000 feet, 2 October 1934, Darlington (MCZ).
Discussion. - Males vary in length from 2.0 to 2.3 mm. and in width from
0.7 to 0.8 mm.; females vary in length from 2.4 to 3.0 mm. and in width from
0.9to1.1mm. Variation in color and pattern is moderate, the minimum
marking consisting of the median thoracic vitta only, the maximum markings
as in the holotype plus a dark spot at the apical termination of elytral intervals
0 and 6, a short vitta on interval 4 before the declivity, and the sutural inter-
val dark on the declivity. The holotype represents the usual color and mark-
ings. Color in one specimen is entirely without green and in another is en-
tirely golden green dorsally. Sculpture of the beak is quite uniform with the
median line more finely marked in females than in males and the interantennal
line usually not marked by even a small glabrous area as in the holotype. The
number of teeth on the fore tibia varies from three to six, with five being the
usual number. The contour of the elytra in profile is not uniform within the
?
36 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
sexes, but the dorsal outline is uniform; the apex is always attenuate in females
and always broadly rounded in males. The setae on the disc of the elytra seem
particularly subject to abrasion, those of the declivity much less so.
The combination of erect elytral setae and the shape of the elytra in dorsal
outline will separate this species from any other tanymecine known from the
West Indies. The other two species in this genus have a pale median vitta on
the pronotum where calypso has a dark median vitta. P. calypso also has
more strongly and more abruptly swollen fore femora than the other Paululusus
and the erect setae are much shorter in constanzae.,
2. Paululusus hispaniolae, new species
Figures 28, 51, 52, 53, 70, 72.
Diagnosis. - Sutural interval swollen at summit of declivity, declivity con-
spicuous in profile. Usually with a pale, median, thoracic vitta.
Description. - Holotype, male, length 2.7 mm., width 1.0 mm. Color
whitish and light brown marked as follows. Sides of beak, head, thorax and
elytra whitish; pronotum with a pale median vitta bordered on either side by a
broad, brown vitta; elytra with a complex pattern of short vittae on the disc and
declivity. Legs feebly mottled; scales of ventrum opalescent or whitish. Scales
of dorsum contiguous, moderate or small, of rounded shapes, granular, feebly
pustulate, with weak to moderate reflexed margins. Setae of head, pronotum
and elytra rather inconspicuous, except in profile, about as long as one scale,
completely arched, uniserial on elytra.
In profile, head subspherical; beak rather short, stout, feebly arcuate.
Dorsal surface of beak (Fig. 28) as wide as head between eyes, sides parallel
except for a feeble emargination of the dorsal-lateral edge before head. Beak
feebly, transversely convex over insertion of antennae, interantennal line
otherwise unmarked; median line marked by a very fine impressed line from
middle of eyes ending in a small but distinct fovea at beginning of interantennal
convexity. Apical emargination acutely angled, occupying approximately one-
half the width of the beak apically, marked posteriorly by a fine carina which
extends caudad from apex for a very short distance. Anterior edge of epistoma
with a median indentation. Scrobe obtusely angled, the horizontal portion
slightly sinuate, the vertical portion curved and strongly tapered posteriorly,
ending beneath anterior edge of eye well above ventral surface; vertical por-
tion about equidistant between eye and apex of beak. Antenna with scape nearly
straight, distal third swollen; funicle with first segment elongate, second seg-
ment one-half the length of the first; segments 3 to 7 subequal, subcubical, the
distal segments broader than the basal ones; antennal club broadly elliptical.
Eye rather large and only moderately convex.
Prothorax 1.1 times longer than wide, sides gently rounded between con-
strictions. In profile, pronotum slightly convex, pronotum 1.5 times longer
than prosternum. Surface of pronotum between constrictions with small, deep
foveae separated from each other by one scale.
Elytra 2.1 times longer than prothorax, elytra across humeri 1.3 times
wider than prothorax. Elytra in dorsal view (Fig. 53) with humeri right-angled,
base obsoletely bisinuate, sides very weakly divergent from humeri to apical
third, thence gently rounded to apex, the apical termination of intervals 4 to 7
protruding slightly into the outline. Elytra in cross-section quite convex; ely-
tra in profile feebly arcuate to summit of declivity. Declivity rounded at sum-
mit, thence slightly concave, the apex of the elytra extending slightly beyond
Howden: West Indian Tanymechini 37
summit; epipleurae slightly emarginate opposite hind coxae, stria 10 obsolete
here. Elytral striae distinct, set with moderate punctures which are separated
from each other in the striae by one scale. Sutural interval at base as wide as
one scale, gradually becoming broader posteriorly; at summit of declivity
rather suddenly swollen and covered with four rows of circular, convex scales.
Remaining elytral intervals slightly convex, subequal.
Fore and hind legs subequal in length, middle leg slightly shorter. Fore
leg with femur moderately swollen, the swelling gradual basally, abrupt distal-
ly; fore tibia straight, slender, with three or four equidistant teeth on inner
edge.
Fore coxae separated by a distance equal to approximately one-third the
width of the antennal club. Anterior portion of ventrites 3 and 4 very narrowly
glabrous across their entire width, slightly concave medially. Ventrites 2, 3
and 4 with their posterior margins slightly convex. Ventrite 5 slightly convex,
its apex without scales and set with numerous fine hairs, its apical margin
truncate and slightly emarginate medially.
Aedeagus almost as long as entire abdomen measured medially. Aedeagus
slender, slightly wider than thickest part of scape; clyindrical; evenly, rather
strongly arcuate; apex in lateral view at a 45° angle; apical opening in dorsal
view elongate oval.
Allotype, female, length 2.9 mm., width 1.2 mm. Differs from holotype
in the following respects. Color pattern more vivid and with fewer pustulate
scales. Prothorax 1.5 times longer than wide; pronotum 1.6 times longer than
prosternum. Elytra much more robust, 2.3 times longer than prothorax; ely-
tra across humeri 1.45 times wider than pronotum, in dorsal view sides dis-
tinctly divergent from humeri to middle thence broadly rounded to declivity,
apex a little narrower and slightly more attenuate. Apices individually rounded.
Elytra in profile (Fig. 51) more arcuate on disc, declivity with sutural interval
at summit more swollen and apex of elytra more attenuate; remaining elytral
intervals flatter than in holotype. Fore coxae not appreciably more widely
separated than in holotype. Ventrite 5 slightly convex along median line with
a weak depression either side near base; covered with scales which are sel-
dom contiguous; apex broadly rounded.
Type series. - Holotype, male, Port au Prince, Haiti, 1 March 1908,
Dr. M. Cameron, B. M. 1936-555 (BM). Allotype, female, same data as
holotype (BM). Paratypes, 22 males, 29 females. DOMINICAN REPUBLIC:
3 males, 4 females, Barahona, September 1938, Darlington (MCZ, Howden);
2 males, 3 females, San José de las Matas, 1000 to 2000 feet, June 1938,
Darlington (MCZ, Howden). HAITI: 1 male, Damien, 8 November 1930, H.
L. Dozier Collector, on Calmite, B. M. 1948-212 (Howden); 1 male, Damiens,
Port au Prince, 9 September 1959, A. M. Nadler (AMNH); 1 male, 2 females,
Gonaives, 4-9 February 1908, Dr. M. Cameron, B. M. 1936-555 (BM, How-
den); 1 female, Jean Rabel, February 1929, E. C. and A. M. Leonard Col-
lectors (USNM); 2 females, Manville, September 1926, Acc. 44-26, G. W.
Wolcott Collector, B. M. 1948-212 (BM); 8 males, 6 females, 32 Km. SW
Mirebalais, c500 feet, 5 June 1956, B. and B. Valentine, beating branches in
semi-desert (Valentine, Howden); 4 males, 9 females, Port au Prince, same
data as holotype (BM, Howden); 1 female, Port au Prince, R. J. Crew, Wick-
ham Collector, 1933 (USNM); 1 female, Port au Prince, 12 December 1928,
Acc. 319-28, A. Audant Collector (USNM); 1 male, Port de Paix, December
1928, E. C. and A. M. Leonard Collectors, on flowering plants (USNM); 1
male, Poste Terre Rouge, 2000 feet, 5 October 1934, Darlington (MCZ).
Discussion. - Males vary in length from 2.3 to 2.9 mm. and in width
from 0.8 to 1.1 mm.; females vary in length from 2.6 to 3.2 mm. and in
width from 1.0 to 1.3 mm. Variation in the series is great, even consider-
38 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
ing the wide distribution on Hispaniola. The aedeagus is particularly vari-
able, but its extreme conditions cannot be correlated with other characters
or with localities. In one extreme (Fig. 70) the apex of the aedeagus is
simple; in the other extreme (Fig. 72) the base of the apex has a flange, a
‘‘collar’’, on either side. The simplest apex is found in specimens from
Port au Prince, the type locality (see Map 4); the most strongly-developed
collar is found in all specimens from San José de las Matas. However,
specimens from Barahona, the extreme southeastern part of the range, have
either a strong collar or an intermediate condition; likewise, the males from
Gonaives and Port de Paix, at the northwestern extreme, have well-developed
collars. Other specimens including the holotype exhibit various intermediate
conditions. | .
Color and color pattern are highly variable, often giving specimens a very
different appearance, but this is not unusual in New World Tanymecini. The
allotype represents the maximum expression of the rather elaborate tan and
white pattern. There are paratypes as well marked, but more often the pattern
is considerably reduced to the vittate prothorax and a pale elytral spot on inter-
vals 5 and 6 at the basal third. There are paratypes from most localities which
are completely pastel green, blue-green or gray-green; in some of these, the
scales, particularly at the base of the elytra, are incompletely developed. In
a few specimens the scales are not margined. The angle of the apical emargina-
tion of the beak varies from right-angled to rather broadly obtusely angled, in
the latter case occupying proportionately more of the width of the beak. The
declivity varies moderately; Fig. 51 (allotype) represents close to the maximum
development, Fig. 52 the weakest. The fore tibiae usually have five teeth on
the inner edge and vary from three (as in the holotype) to six. The separation
of the fore coxae is highly variable, the minimum being as in the holotype, the
maximum being about two-thirds as wide as the antennal club as in some fe-
males.
The swollen sutural interval at the summit of the declivity is quite uniform
in the series and will by itself distinguish this species from all other Paululusus.
3. Paululusus constanzae, new species
Figures 25, 26, 74.
Diagnosis. - Humeri absent, elytra fused and wings absent. Elytral inter-
vals each with a row of erect setae.
Description. - Holotype, male, length 2.3 mm., width 0.8 mm. Dorsum
and legs clothed with whitish and light brown scales arranged as follows: head
pale except for a short brown median vitta on occiput; pronotum with a broad,
somewhat irregular, pale median vitta bordered on either side by a broad,
brown vitta; sides of prothorax mostly pale; elytra indistinctly marked but su-
tural interval on disc pale, base of intervals 6, 7 and 8 pure white, an obscure
‘‘v’? of dark spots beginning about middle and extending to summit of declivity.
Beak clothed with well-separated, smaller, rounded, smooth, submetallic
greenish scales. Ventrum with smooth, shining, opalescent or submetallic
greenish scales. Scales of dorsum and legs not contiguous (except humeral
white spot and a few other dense white areas), finely granular, obsoletely
margined, of irregular polygonal shapes; scales of declivity smaller, more
rounded, smoother. Elytral intervals each set with a single row of erect,
moderately long, slightly arcuate setae, those on disc separated from each
other by three or more scales, those on declivity much more closely placed.
Howden: West Indian Tanymechini 39
Setae of pronotum and occiput very fine, inconspicuous, prostrate; setae of
frons and pronotum apicad of apical constriction thicker, more conspicuous
(especially in profile), more numerous and completely or incompletely arched.
In profile, head subspherical, beak rather thin, its dorsum broadly arcuate.
Head and beak (Fig. 26) with median line impressed from between eyes to inter-
antennal line whence it is feebly carinate to apical emargination. Dorsum with
a vague ‘‘Y’’-shaped depression extending along median line and either side of
apical emargination, the sides of beak convex over scrobes thus creating the
arcuate profile. Beak slender, but by actual measurement wider over scrobes
than head between the eyes; sides of beak conspicuously emarginate before
head. Apical emargination small, occupying less than one-third the width of
the beak anteriorly, acutely angled, marked posteriorly by a very weak carina.
Epistoma with its apical edge irregular, truncate and with several scales en-
croaching upon it. Scrobe obtusely angled, the vertical portion slightly closer
to eye than to apex of beak, ending just below ventral surface of eye. Antenna
with scape nearly straight, distal third swollen; funicle with first segment coni-
cal, second segment elongate-oval and one-half the length of the first segment,
segments 3 to 7 subspherical, distal segments larger; club elongate-elliptical.
Eye moderate in size, moderately convex.
Prothorax 1.2 times longer than wide, sides moderately rounded between
constrictions. In profile, pronotum convex, 1.8 times longer than prosternum.
Surface of pronotum between constrictions with small punctures separated
from each other by a scale.
Elytra (Fig. 25) 1.9 times longer than prothorax; elytra across humeri 0.9
times as wide as prothorax. Elytra fused, wings absent. Elytra narrow,
elliptical, very slightly constricted at apical eighth with apex truncate. Base
of elytra arcuate, humeri absent. Elytra in profile with disc nearly flat to
apical third, thence gently, evenly arcuate to apex. Striae and intervals very
poorly defined; strial punctures small and inconspicuous, their positions ob-
scured by the arrangement of the scales; intervals discernible chiefly by the
rows of setae, intervals on disc as wide as one to two scales, becoming as
wide as two to four scales beyond third interval, especially apically.
Legs long and slender; fore leg conspicuously longer than hind leg, middle
leg conspicuously shorter than hind leg. Fore femur moderately swollen, the
swelling very gradual; fore tibia very slightly sinuate, its inner edge with four
teeth and several denticles all situated on distal two-thirds.
Fore coxae separated by a distance equal to approximately one-third the
width of antennal club. Anterior portion of ventrites 3, 4 and 5 scarcely
modified, very narrowly glabrous and flattened across their entire width. Ven-
trite 5 flattened, slightly convex apically, its surface with only one scale and
rather sparsely clothed with fine hairs of moderate length. Ventrite 5 as wide
basally as long, the sides converging to apex which is broadly truncate and
feebly emarginate.
Aedeagus (Fig. 74) about five-sixths as long as entire abdomen. Aedeagus
very slender, as wide as or slightly wider than thickest part of scape; in pro-
file arcuate, the basal two-fifths more strongly so; apical opening in profile
approximately a 45° angle, in dorsal view elongate oval.
Type series. - Holotype, male, Constanza, Dominican Republic, August
1938, 3-4000 feet, Darlington (MCZ). Paratypes, 4 males: same data as
holotype (MCZ, Howden).
Discussion. - The paratypes vary in length from 2.4 to 2.6 mm. and in
width from 0.8 to 1.0 mm. Two paratypes are colored as the holotype; the
white humeral spot and white sutural interval are constant and characteristic
40 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
of the series. The beak appears to be subject to a moderate amount of varia-
tion; it is usually wider over the scrobes than between the eyes, but in one
specimen is not. In two paratypes the beak is less concave than in the holotype;
in one paratype it is more concave. The apex of the beak, i.e., apicad of the
insertion of the antennae, varies from slightly to strongly deflected. In all the
paratypes the epistoma is wider than in the holotype; one paratype has several
scales encroaching upon the epistoma as in the holotype. The usual number of
tibial teeth is four, but there may be up to six teeth.
When females of this species are found, they may have some development
of the humeral angles, much broader elytra, and the elytra may be longer in
proportion to the length of the prothorax; at least these are the female modifi-
cations exhibited in other flightless Tanymecini such as Pandeleteius dentipes
and Isodacrys spp. There may also be a Slight swelling at the summit of the
declivity, and the elytral striae may be even more confused.
This is the only flightless tanymecine known from the West Indies, the wing-
less condition in this case being considered of specific rather than generic
rank (see discussion following generic description and in the Introduction).
From other species of Paululusus, constanzae may also be distinguished by its
elongate-oval apex of aedeagus and poorly defined elytral striae with erect
setae.
Paradacrys, new genus
Size small. Body and legs covered with scales and small setae. Beak
short, subcubical, its dorsum a little narrowed; apical emargination triangular,
very poorly defined; interantennal line not marked. Scrobe deep, approximately
right-angled; the vertical portion a little closer to eye than to apex of beak, end-
ing before ventral surface. Antenna short, sparsely clothed with very fine
hairs and no scales; scape reaching beyond middle of eye; funicle seven-
segmented, first segment elongate, remaining segments subequal, each ap-
proximately one-half the length of the first and slightly moniliform; club short
to moderate. Eye small, not close to dorsal surface. Prothorax nearly as
wide as long, basal and apical constrictions weak. Pronotum produced anterior-
ly over head, much longer than prosternum; apical margin of prothorax without
ocular vibrissae, teeth or lobes. Elytra with humeral angles well-developed;
10 complete, punctate striae, though stria 10 poorly marked apicad of hind
coxae. Legs short, fore leg only very slightly longer in both sexes. Fore femur
less tapered basally than other femora but scarcely swollen, without teeth or
serrations on inner edge; fore tibia dentate. Fore coxae usually separated by
a width equal to or greater than antennal club. Hind tibia with corbel open.
Tarsi completely padded; tarsal claws free. Abdomen with the anterior portion
of ventrites 3, 4 and 5 slightly modified: a slight depression entire width of
ventrite, but glabrous medially only. Aedeagus slender, as long as first three
and one-half to four and one-half ventrites measured medially.
Type-species. - Paradacrys spatulatum, new species.
The name Paradacrys is intended to mean ‘‘alongside Isodacrys’’ and is
neuter.
Discussion. - Paradacrys is most closely related to Paululusus but it also
shows an affinity with Isodacrys and to a lesser extent with Isodrusus and
Pandeleteinus. Paradacrys is endemic to the Bahama Islands, so it is not
surprising to find its closest relative, Paululusus, endemic to Hispaniola. The
shorter, stouter aedeagus of Paradacrys, its eyes more distant froin the dor-
sal surface, and its much more widely separated fore coxae are considered to
Howden: West Indian Tanymechini 41
be of generic rank and are the principal basis for separating Paradacrys from
Paululusus.
A caricature of Paradacrys might be: an Isodacrys with well-developed
humeri. Less conspicuous distinctions from Isodacrys are: fore coxae a little
more widely separated than is usual for Isodacrys (except I. mexicanum Sharp),
ventrites with weaker modification, aedeagus more slender, scape and funicle
not squamose. Isodacrys is known only from the continental Neotropics (Map
Ty,
Paradacrys differs from Isodrusus in its free tarsal claws, larger beak
and consequently more-oblique scrobe, and much simpler ventrites. I. insul-
anus Howden also occurs in the Bahama Islands.
Paradacrys might also be confused with Pandeleteinus though Pandeleteinus
is not known to occur in the West Indies. Pandeleteinus differs principally in
its contiguous or very narrowly-separated fore coxae, its much broader beak
which is deeply, triangularly excised on the sides, and, usually, its ocular
vibrissae.
The lack of ocular vibrissae in Paradacrys is not emphasized because of
the special situation existing in the West Indies (see Introduction). However,
it is interesting that both species of Paradacrys have a brief row of three or
four very Slightly elongated scales (Figs. 30, 34) on the anterior edge of the
pronotum where ocular vibrissae would occur.
The articulating surface of all tibiae is small and squamose in both species;
however, this may be the consequence of two closely related species rather
than a generic character. The suture separating ventrites 1 and 2 is obsolete
on its median third in both species of Paradacrys; this character may be of
generic rank.
Key to the Species of Paradacrys
1, Elytral setae short, spatulate, i.e., broad and convex at apex.
Scales of elytra small, not contiguous. Aedeagus straight, as long as
first three and one-half ventrites measured medially. .........
ake nt ao tee Geonames Mee 1. spatulatum, new species (p. 41)
Elytral setae longer, ensiform, i.e., narrow, taperedfrom base. Elytra
with white scales, at least, imbricate. Aedeagus slightly arcuate, longer
than tirst four ventrites measured medially. 2.00 5.8 2o).4.52 9 Wo
ee aren aie a en MRM ac peg na ecg Lie 2. ensiformis, new species (p. 43)
(1) Paradacrys spatulatum new species
Figures 31, 34, 35, 46, 55, 73
Diagnosis. - Elytra gently, nearly evenly arcuate from base to apex.
Scales of dorsum small, not contiguous. Elytral setae short, much broader at
apex than at base.
Description. - Holotype, male, length 2.4 mm., width 1.0 mm. Specimen
in poor condition with both antennae mounted separately. Color white and pale
brown, marked as follows. Ventrum and sides of entire body white; an indis-
tinct, median, white vitta on pronotum; elytra indistinctly marked, the white
of the epipleurae extending in an arc onto dorsum from humeri to stria 3 at
about middle, thence receding to stria 4, continuing posteriorly to declivity,
thence across to suture. Scales of dorsal surface (Fig. 46) small, mostly not
42 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
contiguous, weakly margined and alutaceous. Setae of dorsum (Fig. 46 a, b)
shorter than one scale, spatulate, i.e., broader and convex at apex, erect,
feebly arched, uniserial on elytral intervals. Setae more numerous on beak
and becoming longer towards apex of beak. Setae of legs and ventrum longer
and more numerous.
Head and beak in profile flattened from apex to beyond middle of eyes,
thence gently arcuate to pronotum. Beak (Fig. 31) short, subcubical, sides
slightly convergent towards apex. Dorsal surface of beak with a shallow con-
cavity delimited laterally by a faint, obliquely directed ridge extending from
over insertion of antenna to frons; median line deeply impressed on frons; no
interantennal modification. Apex of beak with triangular emargination marked
by a very weak carina and occupying approximately one-half the width of beak.
Epistoma with its anterior edge produced slightly beyond the apex of beak and
reflexed except for small median indentation which is depressed. Scrobe ob-
tusely angled, the horizontal portion shorter and more deeply impressed than
the vertical portion which is slightly curved posteriorly, ending beneath eye
before the ventral surface; vertical portion partly squamose and much closer
to eye than to apex of beak. Antenna short, sparsely clothed with fine hairs.
Scape feebly bowed to fit contour of beak; antennal club short, elliptical, com-
pact. Eye moderate in size, moderately convex.
Prothorax as long as wide, sides feebly protruding, subparallel between
weak basal and apical constrictions; disc flattened. Pronotum 2.0 times
longer than prosternum. Punctures of prothorax obsolete. Anterior margin of
prothorax with no trace of ocular vibrissae; some elongation of scales here
but some scales abraded.
Elytra 2.3 times longer than prothorax; elytra across humeri 1.6 times
wider than prothorax. Elytra with sides subparallel, slightly wider at middle.
Base of elytra slightly arcuately emarginate between fifth striae. Elytra in
profile (Fig. 34) gently, nearly evenly arcuate from base to apex, declivity
only very slightly developed. Strial punctures not very deep and nearly con-
cealed by the vestiture; not precisely aligned throughout; separated in the striae
by one scale. Intervals even and flat, as wide as two to three scales.
Legs short. Fore leg (Fig. 55) scarcely enlarged, only the tibia longer
than that of other legs; fore femur thicker throughout than other femora, but
only slightly thicker medially than at ends; inner edge of fore tibia with four
small teeth and a few minute serrations. Articular surface of tibiae small and
clothed with a few scales.
Fore coxae separated by a distance almost equal to the width of the antennal
club. Abdomen with suture separating ventrites 1 and 2 obsolete on median
third. Anterior portion of ventrites 3 and 4 scarcely modified; anterior portion
of ventrite 5 with median half slightly transversely sulcate. Posterior mar-
gins of ventrites 2, 3 and 4 convex; squamose posterior edge of ventrite 2
touching the succeeding segment throughout, but ventrites 3 and 4 separated
medially from their succeeding ventrites by an abrupt, perpendicular, glabrous,
plane surface, the base of which is slightly anteriorly directed. Ventrite 5
convex, its apex broadly truncate.
Aedeagus (Fig. 73) nearly straight, only basal fourth deflected; slightly
thinner than antennal club; aedeagus as long as length of first three and one-
half ventrites measured medially; apical opening approximately a 45° angle;
apex in dorsal view scarcely attenuate, truncate.
Allotype, female, length 2.8 mm., width 1.3 mm. Color and pattern of
holotype, but form more robust, head and beak stouter, antennal club wider.
Differs from holotype in the following respects. Sculpture of scales a little
Howden: West Indian Tanymechini 43
more pronounced. Dorsal surface of beak much flatter, median impressed
line obsolete on frons, obsolete oblique ridges even weaker than in holotype.
Anterior margin of prothorax at the usual site of ocular vibrissae with a row
of white scales like those of the sides but elongated anteriorly over the mar-
gin. Elytra 2.6 times longer than prothorax; elytra across humeri 1.4 times
wider than prothorax. Elytra with sides subparallel for about basal fifth before
diverging to middle, thence gradually converging, scarcely rounded to apex.
Fore coxae separated by a distance very slightly greater than width of antennal
club. Abdomen (Fig. 35) with ventrites 2, 3 and 4 with posterior margins
scarcely convex, separated from succeeding ventrites almost to sides by an
abrupt, glabrous, plane surface. Anterior portion of ventrites 3, 4 and 5
narrowly glabrous and slightly depressed for approximately median two-thirds.
Ventrite 5 slightly convex, its apex broadly rounded.
Type series. - Holotype, male, Grand Turk, Turks and Caicos Islands,
B.W.I., June 1957, T. H. Farr (CNC). Allotype, female, same data as holo-
type (Inst. Jam.). Paratypes, 3females. BAHAMA ISLANDS: 3 females,
Green Cay (Great Bahama Bank), 18-19 March 1965, B. D. Valentine, R. W.
Hamilton Collectors (AMNH, Valentine, Howden).
Discussion. - The three paratypes vary little fromthe holotype and allotype,
especially in view of the relatively great distance between Grand Turk and
Green Cay. The paratypes range in length from 3.0 to 3.4 mm. and in width
from 1.3 to1.5 mm. Incolor, they vary from whiter and less maculate than
the holotype to much more vividly marked. In the latter, the white elytral
markings are bordered with brown and the disc between the markings is mottled.
White scales tend to be closer to each other than dark scales, but even the
white scales (with rare exceptions) are distinctly separated on the dorsal sur-
face. The dorsum of the beak may be slightly less concave and in one paratype
the frons is much more abruptly transversely swollen. In two paratypes the
elytra are more broadly rounded in dorsal view, the profile is less arcuate,
though not as flat as in ensiformis, and the alternate intervals are slightly
raised, with more numerous setae, and distinctly wider, often being as wide
as six scales. The sutural interval is more elevated for its basal half than in
the holotype or allotype, thus contributing to the arcuate elytral profile.
Paradacrys spatulatum is easily distinguished by its characteristic elytral
setae which are shorter than the surrounding scales, curved and much broader
at the apex than the base; its elytral scales which are very small and not con-
tiguous; its elytral profile which is evenly arcuate from the base; and its aedea-
gus which is straight except at the extreme base.
2. Paradacrys ensiformis, new species
Figures 30, 37, 45, 71.
Diagnosis. - White scales of elytra imbricate, setae of elytra ensiform.
Description. - Holotype, male, length 2.4 mm., width 1.0 mm. Color
white, brown and tan, marked as follows. Head slightly mottled, disc of
prothorax indistinctly marked with three tan vittae. Elytra with epipleurae
and sides of disc white, slightly mottled with tan, the white area reaching stria
6 for basal third, then abruptly swelling in a large lobe to stria 4 at middle of
elytra, receding again to stria 6, then extending diagonally with several inter-
ruptions to summit of declivity; much of the white area bordered anteriorly
with brown; interval 4 brown on basal fourth, remainder of disc mottled tan.
Legs predominantly white; ventrum white. Scales of head and pronotum angular,
44 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
not or almost contiguous, very finely margined, very finely granular. Colored
scales of elytra (Fig. 45a) seldom angular, usually partly or completely
rounded, contiguous or nearly so, very finely margined, very finely granular,
often with an obsolete central pustule, becoming more strongly sculptured to-
wards declivity; most white scales of elytra imbricate. Setae of head and pro-
notum as long as one to one and one-half scales, very slender, nearly prostrate,
slightly arched, numerous on frons; setae on apex of pronotum much longer,
darker. Setae of elytra (Fig. 45a, b) semi-erect, i.e., base of seta arcuate,
remaining half or two-thirds straight, oblique or parallel to surface, ensi-
form, i.e., long, narrow, taperedfrom base. Setae of legs and ventrum like
those of elytra but longer, more conspicuous, more numerous, white.
In profile, beak nearly flat; gradually, slightly deflected apicad of insertion
of antennae; head gently arcuate from posterior edge of eye to pronotum. Dor-
sum of beak 0.9 times as wide as head between eyes, edges straight. Base of
beak with a small, shallow concavity; interantennal line unmarked; scales of
beak gradually becoming more convex, rounded, shinier towards apex, only
slightly sparser; setae of beak straight, erect. Apical emargination poorly
defined, approximately right-angled, its posterior edge not carinate, vibrissae
on sides of emargination indistinguishable from long setae. Median line finely
indented on frons to middle of eyes. In dorsal view, head appears unusually
tapered from base to apex, the posterior edges of the eyes therefore much more
widely separated than the anterior edges. Scrobe slightly obtusely angled,
approximately the same width throughout, deep, horizontal portion three-fifths
as long as vertical portion, slightly closer to eye than to apex, ending above
ventral surface beneath anterior edge of eye. Scape feebly bowed to fit contour
of beak. Antennal club elongate, elliptical, slightly loose. Eye moderate in
size, moderately convex, slightly elliptical.
Prothorax 1.1 times longer than wide; constrictions obsolete on disc, basal
constriction obsolete on sides in dorsal view; sides scarcely rounded between
constrictions. Pronotum 2.0 times longer than prosternum. Punctures of
prothorax obsolete. Anterior edge of pronotum with three consecutive, very
slightly elongated scales at the site of ocular vibrissae.
Elytra 2.3 times longer than prothorax; elytra across humeri 1.3 times
wider than prothorax. Base of elytra slightly arcuately emarginate. Elytra
with sides parallel for basal fourth, feebly divergent to just beyond middle,
thence gradually, broadly rounded to declivity, the apex broadly rounded, ex-
tending slightly beyond; apices briefly, slightly divergent along suture; indivi-
dually rounded. In profile, elytra flattened on basal fourth, slightly arcuate,
thence more strongly arcuate from basal third; summit of declivity scarcely
perceptible. Strial punctures moderately small, regularly aligned. Intervals
even, very slightly convex on disc.
Fore leg distinctly longer than other legs, fore femur proportionately not
thicker than other femora; inner edge of fore tibia with four moderate teeth.
Articular surface of tibiae densely, almost completely squamose.
Fore coxae separated by a distance equal to four-fifths the width of the
antennal club. Abdomen (similar to Fig. 37) with suture separating ventrites
1 and 2 obsolete on median third. Ventrites 3, 4 and 5 with anterior portion
glabrous and concave for approximately median three-fifths, the modification
wider medially; edge of modification not abrupt, partly squamose. Posterior
edge of ventrites 2, 3 and 4 abruptly perpendicular, glabrous opposite modifi-
cation of anterior edge of succeeding segment. Ventrite 5 nearly flat, apex
deflected, briefly truncate-emarginate.
Aedeagus (Fig. 71) slightly longer than first four ventrites measured
Howden: West Indian Tanymechini 45
medially; slightly arcuate, basal fourth more abruptly so; thickened before
apical opening which is oval with apex attenuate. Aedeagus thicker than an-
tennal scape, but thinner than antennal club.
Allotype, female, length 3.1 mm., width 1.3 mm. A fine, fresh specimen
with mandibular cusps still attached (Fig. 30). Differs from holotype in the
following respects. Color pattern as in holotype but thoracic markings less
distinct on disc and elytra darker on disc. More elytral scales imbricate than
in holotype. Head and beak more robust; eyes less convex, scarcely exceeding
sides of beak in anterior view. Median line impressed for distance of only one
scale. Scrobe with horizontal portion three-fourths as long as vertical portion;
antennal club slightly shorter, broader than that of holotype. Elytra 2.4 times
longer than prothorax; elytra across humeri 1.4 times wider than prothorax.
Elytra with sides more gradually converging from middle, apex narrower,
longer; in profile, declivity more oblique. Alternate intervals very slightly
wider, intervals across middle of elytra as wide as three to five scales. Fore
coxae separated by distance approximately equal to width of antennal club.
Modification of anterior portion of ventrites 3 and 4 similar to holotype; modi-
fication of ventrite 5 (Fig. 37) much broader, approximately rectangular,
occupying approximately median half of ventrite. Ventrite 5 elongate, very
narrowly rounded.
Type series. - Holotype, male, 5 miles E. lighthouse, S. shore, Castle
Is., Bahama Islands, B.W.I., 4 April 1965, B. D. Valentine, R. W. Hamilton
Collectors (AMNH, temporarily Valentine). Allotype, female, same data as
holotype (AMNH, temporarily Valentine). No paratypes.
Discussion. - Paradacrys ensiformis may always be distinguished from
P. spatulatum by any of the following characters: aedeagus long, arcuate;
many scales of elytra imbricate; setae long, slender, semi-erect. The less
evenly arcuate elytral profile is also distinctive but more difficult to assess.
The type locality of P. ensiformis is nearly in the middle of the rather
large range of P. spatulatum.
Pandeleteius Sch6nherr
Pandeleteius is a very large New World genus but is represented in the
West Indies by only two species: the endemic Pandeleteius testaceipes Hustache
and the introduced Pandeleteius nodifer Champion.
In this genus the anterior portion of ventrites 3, 4 and 5 (Fig. 44c) is flat
and unmodified (rarely arcuately depressed and partly glabrous) though the
posterior portion may be modified; the fore legs are usually conspicuously
enlarged; and the ocular vibrissae are well-developed.
Key to the West Indian Species of Pandeleteius
1. Immaculate or predominantly white. Elytra with base of intervals 3, 4
and 5 conjointly produced anteriorly. ... 1. nodifer Champion (p. 46)
Maculate brown? Base of elytra unmodifieds sy s0 oo ee es I a,
Dia veer Oe ile Pee PER GRR REP SE “sh CRA ELEN aR 2. testaceipes Hustache (p. 48)
46 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
1. Pandeleteius nodifer Champion
Figures 44c, 76.
Pandeleteius nodifer Champion, 1911, p. 206. Lectotype, here designated,
female, labelled, ‘‘Lago Managua, Nicaragua, 7.II. Solari’’, ‘‘Type’’
ona circle of paper with an orange border, ‘‘9’’, ‘‘Sp. figured’’, and
‘‘vibr. wanting’’ (BM).
Diagnosis. - Elytra with base of intervals 3, 4 and 5 conjointly produced
anteriorly.
Description*. - Males vary in length from 4.3 to 5.2 mm., and in width
from 1.3 to1.7 mm. Females vary in length from 4.4 to 5.6 mm. and in
width from 1.6 to 2.1 mm. Clothed with white, marked with a pale brown,
broad, median thoracic vitta and infrequently with scattered spots on elytra.
Scales mostly not contiguous, finely granular, not margined. Setae minute.
Beak flat or feebly concave at base, long; sides parallel, vertical. Median
line impressed from about middle of eyes to interantennal line (which is un-~
marked), from thence median line finely glabrous, carinate to apical emargina-
tion. Apical emargination acutely angled, as long as wide, finely carinate,
occupying less than one-half the width of beak. Epistoma slightly concave,
anterior edge emarginate medially. Scrobe gently arcuate, ending opposite
middle to ventral edge of eye. Eye very large, flattened; scarcely more con-
vex than head.
Prothorax averaging 1.2 times longer than wide in males and 1.1 times longer
than wide infemales. Prothorax widest at apical third (over fore coxae); con-
strictions absent or obsolete on sides. Disc flattened, depressed before base,
leaving base higher than disc. Pronotum 1.1 times longer than prosternum.
A cluster of an average of 12 vibrissae arising from a squamose tubercle,
the vibrissae often reaching eye.
Elytra of males average 2.3 times longer than prothorax, elytra of females
average 2.6 times longer than prothorax; elytra across humeri 1.25 times wider
than prothorax in both sexes. Males with sides of elytra nearly parallel to mid-
dle, thence converging to apex, scarcely constricted beneath terminus of inter-
vals 4 to 6; in profile, disc flat, declivity oblique to nearly vertical; summit of
declivity rounded. Females with sides of elytra rounded, divergent beyond
basal fifth or sixth, convergent from about middle; apical terminus of intervals
4 to 6 somewhat more conspicuous in dorsal outline; in profile, disc slightly
arcuate beyond basal fifth or sixth, sutural interval at summit of declivity pro-
duced posteriorly in a knob, declivity arcuate or straight, slightly oblique.
Base of elytra with intervals 3, 4 and 5 conjointly arcuately produced anteriorly
and upward. Elytral striae closely set with small punctures; striae not always
perfectly straight; striae 1, 2 and 3 in particular prone to irregularities on
declivity. Intervals becoming convex towards sides and base.
Fore leg long; fore femur greatly but gradually swollen; fore tibia slender,
with five to seven small teeth on inner edge.
Fore coxae of male extremely narrowly separated, i.e., by less than
narrowest width of scape; fore coxae of female more widely separated but still
usually less than greatest width of scape. Ventrite 5 of male nearly flat, apex
2k
This description, except for measurements, is based on the Jamaica population;
comparison of non-West Indian populations is found under ‘‘ Discussion” .
Howden: West Indian Tanymechini 47
truncate to emarginate.
Aedeagus (Fig. 76) evenly, rather strongly arcuate; dorsum depressed from
apical third to orifice which is at apical sixth; apex turned up. Ventrite 5 of
females with slight depression on either side at base; apex narrowly rounded.
Type material. - Paralectotype, hereby designated, male, labelled
‘<Colombia’’ (BM).
The lectotype is 4.6 mm. long, 1.7 mm. wide. It is teneral, and missing
the last three tarsal segments on the left fore leg as well as a few scales of
dorsal surface. The color in the illustration is accurate. The lectotype differs
from the description in the following respects: the prothorax instead of being
‘‘as long as broad’’ is actually much longer than broad, i.e., 3.1:2.8. Champi-
on describes the vibrissae as ‘‘reduced to a few hairs’’; actually they are abra-
ded on the right side and on the left side consist of a fully-developed cluster.
Distribution. - Colombia (BM, MCZ), Nicaragua (BM), Honduras (Howden,
Hubbell, USNM), Jamaica (Inst. Jam., Ill.), and Miami, Florida (Fla. Plant
Board, Howden, Kissinger, USNM). Total number of specimens examined:
32. JAMAICA: 4 males, 8 females, Port Henderson Hill, St. Catherine, 25
June 1958, M. W. Sanderson or T. H. Farr, on Cassia emarginata (Inst. Jam.,
Ta.)
Discussion. - In addition to the population on Jamaica, a colony of P. nodi-
fer now exists around Miami, Florida. The Miami specimens differ as follows:
scales often much more closely spaced and in a few specimens are even imbri-
cate; thoracic vitta pale gold, no specimens with elytral spots; up to ten teeth
and many extra serrulations on inner edge of fore tibia.
Honduras specimens, as would be expected, conform most closely to the
lectotype. Compared to the Jamaican specimens their scales are more coarse-
ly granular, in some specimens being even subpustulate and submarginate, and
elytral spots are much more frequent.
In one Colombian female the apex of the beak is less deflected, and in an-
other the color pattern is much more elaborate: intervals 3 and 4 white, inter-
vals 1 and 2 brown and remainder of disc of elytra mottled. Thus the most
immaculate specimens occur in the northern part of the range (Florida) and the
brown markings become more frequent and more numerous southward, being
most fully developed in the southern extreme (Colombia).
There are at least nine species of white Pandeleteius on the continent; of
these at least two - amulae Champion and albisquamis Champion - have some
development of the base of the elytra. Both of these species differ from
nodifer in having the prothorax much shorter and the fore coxae widely separ-
ated. Pandeleteius nodifer is very close to longicollis Champion, another
white species which has a large distribution, but in this case along the Gulf
Coast of Mexico into Texas. Pandeleteius longicollis differs from nodifer in
the lack of development of the base of the elytra and the orifice of the aedeagus
situated at the basal two-fifths. .
Biology. - A variety of hosts are recorded for the adults. Honduras speci-
mens are recorded from leaves of Lantana camara L., carbonal, and from the
‘‘roadside’’. The Jamaican specimens were taken on Cassia emarginata. The
Florida specimens were taken on ‘‘bush’’ and ‘‘feeding on the leaves of
Pithecolobium dulce Benth. ”’
48 Contrib. Amer. nt. Inst.., vol. 5, no..5; 1970
2. Pandeleteius testaceipes Hustache
Figures 36, 75
Pandeleteius testaceipes Hustache, 1929, pp. 181-182. Cotypes, Guadeloupe
(Paris) (See discussion under ‘‘Type Material’’).
Diagnosis. - Posterior margin of epistoma parabolic, strongly and acutely
keeled, extending half-way to interantennal line.
Description. - Males, length 2.5 to 3.1 mm., width 0.9 to 1.1 mm. ; fe-
males, length 3.4 to 3.7 mm., width 1.3 to 1.4 mm. Color and markings as
described by Hustache, but museum specimens brown rather than testaceous.
Brown and pale brown with sides and ventrum pale or opalescent; disc of thor-
ax with a pale vitta either side of median line, elytra sometimes with a broken
white ‘‘V’’? at summit of declivity, disc of elytra usually mottled. Scales finely
granular, not margined or with a margin which is usually composed of coales-
cent granules. Elytral setae very fine, inconspicuous, completely arched,
uniserial on all intervals.
Beak rather elaborately sculptured; sides subparallel from base to apicad
of scrobes, thence abruptly convergent. Dorsal surface of beak with lateral
ridges extending obliquely towards median line, their termination at base of
beak abrupt, often creating a transverse depression between eyes; interantennal
line marked by a fine to moderate ridge; median line moderately to deeply im-
pressed from interantennal ridge to frons; scales apicad of interantennal ridge
usually submetallic. Epistoma extending approximately one-half of the distance
from apex to interantennal line; posterior margin of epistoma parabolic in shape,
strongly and acutely keeled, set with a row of three or four vibrissae; anterior
edge of epistoma straight with a small median indentation. Scrobe deep, broad,
the vertical and horizontal portions approximately equal and at right angles to
each other, ending before ventral surface. Antenna with club usually approxi-
mately twice as long as wide, the segments of the club somewhat ‘‘loose’’.
Eye moderate in size and convexity.
Prothorax usually as long as wide, but in males sometimes longer than wide
and in females sometimes wider than long. Thorax with moderate basal and
apical constrictions, sides rounded between constrictions, disc only feebly con-
vex, median line unmarked except by a narrow dark vitta. Pronotum slightly
produced anteriorly over head; in males pronotum averages 1.4 times longer
than prosternum, in females pronotum averages 1.6 times longer than pro-
sternum. A cluster of four or five ocular vibrissae arising from the unmodi-
fied margin of pronotum.
Males (Fig. 36) with elytra subcylindrical, 2.5 to 2.8 times longer than
pronotum, sides parallel to about apical third, thence gently rounded to beneath
termination of intervals 4 to 7, beyond which the apex is slightly attenuated.
Females with elytra approximately diamond-shaped, 2.8 to 3.0 times longer
than pronotum, sides divergent from humeri to just beyond middle, thence con-
vergent to apex with scarcely any interruption in dorsal outline beneath termina-
tion of intervals 4 to 7; in profile, disc beyond basal fourth much more convex
than in males. Elytral intervals equal, flat or nearly so. Striae set with small
punctures, not very prominent.
Fore femur strongly, abruptly swollen; fore tibia slender, feebly bowed or
straight, usually with six small teeth on inner edge. Tarsal claws free.
Distance between fore coxae of males equals approximately half the width
of antennal club; between fore coxae of females equals the width or slightly more
Howden: West Indian Tanymechini 49
than width of antennal club. Ventrite 5 of male long, slightly convex, with
apex broadly rounded. Ventrite 5 of female triangular in shape with apex
narrowly rounded, a shallow depression each side of median line at base.
Aedeagus (Fig. 75) straight with basal third or fourth and apex curved
downward. In dorsal view apical opening broad, wider than shaft of aedeagus,
elongate elliptical.
Type material. - The type series is listed by Hustache (1929, p. 182) as
consisting of ‘‘15 specimens (Fl. 8; Mus. 17)’’; only the ‘‘Fl. 8’’ have been lo-
cated to date. In the introduction to his paper, Hustache relates that the publi-
cation of Fleutiaux and Sallé’s list of Coleoptera of Guadeloupe (1889) was a
great stimulus to collecting on Guadeloupe, G. Dufau in particular responding
by collecting and sending to Paris approximately 9,500 Curculionidae for deter-
mination! Dufau sent three principal collections - the first to Fleutiaux, the
second to the Museum, and later a third collection to Hustache. In the Paris
Museum I searched unsuccessfully for P. testaceipes in the general collections
and the Hustache collection. Mme. Bons later located the series of eight speci-
mens in Box no. 1 of the Fleutiaux Collection. The series (except ‘‘the type’’)
was very kindly loaned for study and there seems no reason to doubt that they
are the ‘‘Fl. 8’’ of Hustache. However, I doubt that the remaining specimen
is the type, since Vaurie (in litt.) reports that it does not bear a type label,
whereas all the other Hustache species that I saw bear a white label with
‘““TYPE’’ printed in red. This specimen is labelled in Hustache’s handwriting.
‘*Pandeleteius testaceipes mihi’’, and above this label is another, in the same
writing ‘‘P. crassipes, n. sp.’’; it is from Trois Riviéres, Guadeloupe, Dufau
Collection. ‘*P, crassipes’’ could well be a Dufau name which Hustache decided
not to perpetuate.
A lectotype is not designated since the type could well be among the seven
specimens that have not been located; there is no taxonomic confusion between
this species and any others. |
Distribution. - Lesser Antilles. Total number of specimens examined: 19.
DOMINICA: 1 male, Gran Savanna Nr. Salisbury, 18 June 1964, H. Hespen-
heide, on Crotan sp. (Hespenheide). GRENADA: 1 male, Balthazar, Windward
side, H. H. Smith, 1917-125, 38 (BM); 1 male, Windsor, H. H. Smith 25 (BM).
GUADELOUPE: 1 female, Gourbeyre, Vitrac (Fleutiaux Coll., Paris, cotype);
3 males, 2 females, Pointe Noire, Route des Mamelles, 25 August 1965, J.
Bonfils (AMNH, Howden); 3 males, Trois Riviéres, Dufau, one with additional
data of: au parap en forét trés rare, exemplaire foncé (Fleutiaux Coll., Paris,
cotypes); 3 males, Vitrac (Fleutiaux Coll., Paris, cotypes). ST. VINCENT:
2 males, 2 females, H. H. Smith, 1917-125, one male with additional data:
3000 feet, Windward side (BM, Howden).
Discussion. - P. testaceipes is most closely related to Pandeleteius
kirschi (Faust)* which occurs in Venezuela and Colombia. A series of ten
specimens of kirschi, including the type, was examined and compared with
testaceipes. Other than the differences in the range and in size (the mainland
kirschi males average 0.3 mm. longer and the females 0.5 mm. longer),
testaceipes can be separated from kirschi by the: aedeagus which is rather
broadly rounded apically (in kirschi pointed and with a knob at apex), elytra
slightly attenuate in female, epistomal keel extending about half way to inter-
antennal line (in kirschi keel extends three-fourths of the distance to inter-
antennal line), antennal club which is slightly shorter, broader and ‘‘looser’’,
and less swollen fore femora.
To be discussed in a future paper.
50 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
Of all the endemic West Indian Tanymecini, this is the only one which
bears ocular vibrissae. They are only scarcely reduced from the luxuriant
vibrissae of P. kirschi.
The three specimens of Pandeleteius sublineatus Champion from St. Vin-
cent and Grenada which Champion said, ‘‘seem to belong to this species’’
[sublineatus] (1911, p. 204) are actually P. testaceipes. These are also the
specimens referred to sublineatus in Leng and Mutchler’s list (1914, p. 468)
of the Coleoptera of the West Indies.
Isodrusus Sharp
Isodrusus is represented in the West Indies by two endemic species, insul-
anus Howden in the Bahama Islands and guajavus n. sp. in Jamaica. The third
species in the genus, debilis Sharp, occurs on the mainland from Guatemala to
Texas (Map 3). The species are quite uniform in their very short beaks;
strongly modified anterior portion of abdominal ventrites 3, 4 and 5; antennal
scrobe strongly angled and deep to ventral surface of beak; and, of course,
connate tarsal claws.
Key to the West Indian Species of Isodrusus
1. Fore tibia straight, with five to eight teeth on inner edge. Ocular vibrissae
present (but vestigial). Dorsal surface of beak not constricted or nar-
rowed at base. Bahama Islands..... 1. insulanus Howden (p. 50)
Fore tibia strongly bowed, without teeth on inner edge. Ocular vibrissae
absent. Dorsal surface of beak constricted at base to one-half the width
of the head between the eyes. Jamaica. .......6+e6.s8+s808e88080-:
1. Isodrusus insulanus Howden
Isodrusus insulanus Howden, 1963, pp. 43-45. Type, Long Island (USNM).
Diagnosis. - As in key.
Discussion. - No additional specimens have been seen of this species since
it was described from Long Island and Egg Island in the Bahama Islands. The
aedeagus is Similar to that of I. guajavus but it is more evenly arcuate and the
apical opening is broader. The radial pattern of the elongate scales is quite
distinctive. Length ranges from 2.5 to 3.7 mm., considerably larger than
guaj avus.
2. Isodrusus guajavus, new species
Figures 32, 33, 56, 69.
Diagnosis. - Fore tibia strongly bowed, without teeth on inner edge. Dor-
sum of beak greatly reduced in size at base by encroachment of scrobe.
Description. - Holotype, male, length 2.0 mm., width 0.8 mm. Color
light green and bronze, marked as follows. Head and thorax mottled with
sides mostly green; elytra with disc predominantly bronze, sides of elytra
mostly green, the green protruding upon disc in a lobe to stria 4 at basal third
and to stria 2 at apical third. Scales rounded or angular in shape, not arranged
Howden: West Indian Tanymechini aL
radially around punctures, finely margined, reticulate to coarsely granular;
green scales more finely sculptured and seldom margined; scales mostly not
contiguous. Setae small, very inconspicuous, about as long as one scale,
slender, completely arched.
In profile, head and beak obsoletely arcuate to posterior half of eyes, then
feebly arcuate to pronotum. Beak short, cubical, three-fourths as long as
head between eyes; sides vertical. Scrobes convergent towards base of beak,
encroaching upon dorsal surface of beak and greatly reducing its size. Dorsum
of beak at base 0.5 times as wide as head between eyes; dorsum at widest point,
over insertion of antennae, 0.8 times as wide as head between eyes. Median
line impressed from between insertion of antennae to middle of eyes; dorsum
of beak longitudinally convex either side of median line to apicad of insertion
of antennae, thence entire apex of beak abruptly, obliquely deflected. Apical
emargination obliquely angled, its apex rounded, occupying approximately
one-half the width of the beak; margin obsoletely carinate; epistoma with its
anterior edge straight; with three vibrissae on each side. Scrobe acutely
angled, shaped like a ‘‘7’’, vertical portion bowed towards apex of beak, deep
and well-defined throughout, ending on ventral surface in front of anterior edge
of eye. Scape bowed, much more strongly so basally. Antennal club moderate
in size, base slightly elongate, cylindrical before expanding. Eye large,
moderately convex, large-faceted.
Prothorax 1.2 times longer than wide; sides moderately rounded between
relatively wide, equal basal and apical constrictions. Pronotum 2.0 times
longer than prosternum; in profile, disc distinctly arcuate, constrictions dis-
tinct, the apical one longer. No trace of ocular vibrissae. Punctures small
to moderate, deep.
Elytra 2.1 times longer than prothorax; elytra across humeri 1.3 times
wider than prothorax. Base of elytra slightly emarginate. Elytra with sides
parallel for basal seventh; slightly divergent from there to apical three-fifths,
thence convergent, gently rounded to apex, the apical terminus of intervals 4
to 6 very slightly visible in dorsal outline. In profile, (Fig. 32) disc of elytra
nearly flat, not deflected until beyond apical three-fifths; declivity straight,
slightly oblique. Intervals even, flat. Striae set with moderate, deep punctures,
becoming much smaller and shallower towards apex.
Fore leg (Fig. 56) with femur moderately, abruptly swollen. Fore tibia of
even thickness; strongly, evenly bowed; with four minute denticles on inner
edge. :
Fore coxae separated by less than the greatest width of the antennal scape.
Modification of anterior edge of ventrites 3, 4 and 5 almost straight, abrupt
across entire width, higher medially, the edge not carinate. Ventrite 5 moder-
ately convex, its apex narrowly truncate.
Aedeagus (Fig. 69) arcuate with apical half less so, slightly thicker than
the apex of scape, as long as entire abdomen measured medially; apical open-
ing in dorsal view elliptical.
Allotype, female, length 2.1 mm., width 0.8 mm. Differs from holotype
as follows. Pronotum with a brown median vitta; elytral green markings
bordered with brown. Head and beak more robust but dorsum of beak no larger
and hence even more strongly constricted at base in relation to width of head
between eyes. Prothorax 1.03 times longer than wide; pronotum 1.7 times
longer than prosternum. Elytra 2.4 times longer than prothorax; elytra across
humeri 1.3 times wider than prothorax. Elytra in dorsal view wider than in
holotype at widest point, convergent sides less rounded to apex. Declivity
52 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
more abrupt, less oblique, obsoletely concave. Modification of anterior por-
tion of ventrites (Fig. 33) 4 and (particularly) 5 arcuate, high medially, the
edge subcarinate. Ventrite 5 slightly depressed behind anterior modification,
otherwise slightly convex and scarcely distinguishable from that of male.
Type series. - Holotype, male, Moneague, St. Ann, Jamaica, 20 August
1966, A. T. Howden, on guava (Howden). Allotype, female, same data as
holotype (Howden). Paratypes, 41 males, 40 females. JAMAICA: 37 males,
35 females, same data as holotype, A. T. or H. F. Howden collector (CNC,
Howden); 2 males, Ewarton, St. Catherine Parish, 28 September 1945, E. L.
Sleeper Collector (Sleeper); 2 males, 4 females, Newport [Manchester Parish],
18-19 February 1937, Chapin and Blackwelder (USNM); 1 female, Port Hender-
son, St. Catherine Parish, 28 September 1945, E. L. Sleeper Collector
(Sleeper).
Discussion. - Males vary in length from 1.8 to 2.0 mm. and in width from
0.7 to 0.8 mm.; females vary in length from 1.9 to 2.2 mm. and in width from
0.7to 0.9 mm. Variation in the type series is dight. The scales may be
more or less sculptured than in the holotype. The green scales may be re-
placed in part or rarely entirely by white or tan and the bronze by light brown.
The characters of the beak are particularly constant, though the dorsum may
have the longitudinal convexities flatter than in the holotype.
The sexes are difficult to separate externally, the only reliable character
being the anterior modification of ventrite 5 which is distinctly arcuate in the
female and nearly straight in the male.
Isodrusus guajavus may always be distinguished from other Isodrusus by
the conspicuously bowed, non-dentate fore tibiae and the greatly reduced dor-
sal surface of the base of the beak. Isodrusus guajavus is the smallest Iso-
drusus and the only one with green scales. The eye of guajavus is slightly
smaller than that of insulanus, but much larger than that of debilis.
The series from Moneague was collected in a small, hilly pasture on young
guava trees infruit. It was not determined if the beetles were feeding on the
fruit, leaves or stems; their small size and coloration thoroughly camouflaged
them on the hairy trees.
The Ewarton and Port Henderson specimens collected by E. L. Sleeper were
taken on a mesquite-like tree (Acacia?) called Huisache (in litt. ).
FAUNAL LIST BY ISLANDS
Bahama Islands
Castle Island
Paradacrys ensiformis, new species
Egg Island
Isodrusus insulanus Howden
Grand Turk
Paradacrys spatulatum, new species
Green Cay
Paradacrys spatulatum, new species
Long Island
Isodrusus insulanus Howden
Rum Cay
Scalaventer convexifrons, new species
Howden: West Indian Tanymechini 3 Pe
Greater Antilles
Cuba
Scalaventer cubensis, new species
Scalaventer subtropicus (Fall)
Grand Cayman
Scalaventer caymani, new species
Jamaica
Scalaventer cyrillae, new species
Scalaventer coccolobae, new species
Scalaventer litoreus, new species
Scalaventer montanus, new species
Scalaventer remotus, new species
Scalaventer jamaicensis, new species
Scalaventer valkyrius, new species
Isodrusus guajavus, new species
Pandeleteius nodifer Champion
Hispaniola
Scalaventer gelinasus, new species
Paululusus calypso, new species
Paululusus hispaniolae, new species
Paululusus constanzae, new species
Lesser Antilles
Guadeloupe
Pandeleteius testaceipes Hustache
Dominica
Pandeleteius testaceipes Hustache
St. Vincent
Pandeleteius testaceipes Hustache
Grenada
Pandeleteius testaceipes Hustache
ZOOGEOGRAPHIC RELATIONSHIPS
A discussion of zoogeography can be only as sound as the extent to which
the fauna is known, and there is still much to be learned of the West Indian
Tanymecini. However, it seems there has been enough collecting by enough
entomologists to consider the fauna of the islands proportionately represented
and thereby indicate zoogeographic trends.
Here, as in the rest of the paper, the genera Pachnaeus and Polydacrys
are not discussed; they are widespread in the West Indies.
In the West Indies there are five genera of Tanymecini of which two are
truly endemic. By comparison, the total New World fauna is 16 genera (in-
cluding several undescribed genera), all but one (Tanymecus) restricted to
the New World. The relationships of the endemic genera to the mainland forms
can only be surmised by morphological similarities.
Morphologically, the West Indian fauna is relatively unique among New
World tanymecines in the modification of the anterior portion of the abdominal
ventrites 3, 4 and 5. This modification is strong in Scalaventer and Isodrusus,
reduced in Paululusus and greatly reduced in Paradacrys; there is no modifica-
tion in the fifth genus, Pandeleteius. The most extreme development is found
in Scalaventer cyrillae, endemic to the high mountains of Jamaica. Other
tanymecines having the anterior portion of the abdominal ventrites modified
54 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
are: Isodacrys, Pandeleteinus and the Pandeleteius subgenus Exmenetypus
Voss. In this respect and others these taxa and the West Indian endemics are
interrelated. Isodacrys ranges from southwestern United States to Guatemala,
Pandeleteinus ranges from southwestern United States to Guerrero, Mexico;
Exmenetypus ranges from Guatemala to Venezuela. See Maps 1, 2 and 3.
The island Tanymecini readily fall into two groupings, essentially following
the classic pattern of a division between the Greater and Lesser Antilles.
In the Lesser Antilles the sole tanymecine representative is Pandeleteius
testaceipes, which is endemic to Guadeloupe, Dominica, St. Vincent and
Grenada. The species is closely related to Pandeleteius kirschi of Venezuela
and Colombia, and one can easily postulate a South American origin for P.
testaceipes.
The species occurring in the Greater Antilles and the Bahama Islands (see
preceding Faunal List by Island) present a complex picture. The relatively
rich Bahamian fauna consists of three genera, one of them endemic (Paradacrys).
The two species of Paradacrys seem most closely related to Paululusus, ende-
mic to Hispaniola. The other two genera on the Bahamas, Scalaventer and Iso-
drusus, have relatives on the Greater Antilles and the mainland. The presence
of one endemic genus, plus four endemic species, seems indicative of a rela-
tively long separation of the Bahamian fauna.
The tanymecines on the remaining group of islands, Jamaica, Hispaniola,
Cuba and Grand Cayman can be more easily related. Three genera occur in
this area; two of them, Isodrusus and Scalaventer, are each represented on
the mainland by a single species.
Only three species of Isodrusus are known. Isodrusus debilis, occurring
from Texas to Guatemala, has well-developed vibrissae. Since the presence
of ocular vibrissae is essentially a tribal character, the reduction or absence
of vibrissae is significant, complete loss being considered indicative of long
isolation. The Bahamian insulanus has vestigial vibrissae and the Jamaican
guajavus has no vibrissae. This would indicate a longer isolation in Jamaica.
A more specific interpretation based on the three species, other than consider-
ing that their distribution forms a relict pattern, is not feasible. In the case
of Scalaventer, the mainland species, subtropicus, has an unusual distribution,
occurring in Florida, Baja California and Cuba. Specimens from the two
mainland localities have been carefully checked and seem identical. Since the
two specimens known from Baja California are labelled merely ‘‘Santa Rosa,
Low, Calif.’’ with no other data, it seems inadvisable to place too much weight
on the odd distribution. The Florida records represent numerous collections
and a Cuba-Florida distribution is not unusual. The closest relatives of sub-
tropicus are caymani and jamaicensis, and to a lesser degree the Bahamian
convexifrons, the four species forming a distinct group. For this group the
invasion pattern is assumed to stem from a center in Cuba or Florida. Two
other very distinct groups, composed of five species, are endemic to Jamaica,
which indicates a fairly long isolation and probably more than one invasion.
Hispaniola, while having only a single Scalaventer, has three species of
the endemic genus Paululusus. Again there is the inference of a relatively
old, isolated fauna. This coincides with the findings in other groups.
There remains to be discussed Pandeleteius nodifer on Jamaica. I feel
this is relatively recently established on Jamaica since it is widespread in
Central America, is apparently confined to the Port Henderson area in Jamaica,
is also recently established around Miami, Florida, and no other species of
the large genus Pandeleteius occurs in the Greater Antilles.
Apparently, none of these tanymecines have reached Puerto Rico, or do
Howden: West Indian Tanymechini 55
not now occur there. Extensive collecting in Puerto Rico during July, 1969,
yielded no specimens pertinent to this study, none were found in any of the
collections examined and there are no records in the literature. The Lesser
Antillean fauna does not extend as far northward as Puerto Rico, and the
Greater Antillean fauna veers northward and westward to the Caicos and
Bahama Islands, not reaching Puerto Rico.
As far as is known none of the adult food plants in this study are endemic
to the West Indies though most are native.
In summary, the tanymecine genera considered here probably represent
early invasions of the islands, the Greater Antillean fauna either from Yucatan
to Cuba, from Nicaragua-Honduras to Jamaica, or Florida to Cuba and thence
through Hispaniola to Jamaica. The Bahamian genera may have derived from
Floridian stock (especially Scalaventer and Isodrusus) as well as Greater
Antillean stock (Paradacrys). The single Lesser Antillean species derived
from Venezuelan stock. Why all of the genera mentioned are absent from |
Puerto Rico remains an interesting question.
ACKNOWLEDGMENTS
A substantial part of the information in this paper results from a two month
collecting trip to Jamaica which was supported by Grant No. 4145, Penrose
Fund, of the American Philosophical Society. The results of this trip are re-
corded in the Year Book of the Society for 1967. The Institute of Jamaica and
in particular Dr. Tom Farr extended many courtesies in connection with this
trip and their assistance is gratefully acknowledged. Dr. George Proctor also
of the Institute of Jamaica kindly identified the host plants; the voucher speci-
mens are deposited in that collection.
The lectotype of Pandeleteius nodifer was selected while studying at the
British Museum in 1962 on Grant No. 438, Johnson Fund, American Philo-
sophical Society.
Additional specimens of West Indian Tanymecini were borrowed from the
following institutional and personal collections: American Museum of Natural
History (AMNH), Patricia Vaurie; British Museum (Natural History) (BM),
Richard Thompson; Canadian National Collection (CNC); Illinois Natural His-
tory Survey (Ill.), John Kingsolver and Milton Sanderson; Institute of Jamaica
(Inst. Jam.); California State College at Long Beach (Long Beach); Museum of
Comparative Zoology (MCZ), P. J. Darlington, Jr., and John Lawrence;
Muséum National d’Histoire Naturelle (Paris), A. Descarpentries; United
States National Museum (USNM), Rose Ella Warner; Henry A. Hespenheide;
Elbert Sleeper; and Barry D. Valentine.
Numerous facilities of the Entomology Research Institute, Canada Depart-
ment of Agriculture, were available to me and I would like to thank the Institute
for this cooperation. Both my husband, Henry, and Edward C. Becker of the
Institute were especially helpful with several aspects of the study in Ottawa and
in Jamaica.
A search for the type of Pandeleteius testaceipes Hustache involved M.
Jacques Bonfils, Petit-Bourg, Guadeloupe; Mme. Bons, Muséum National de
l’Histoire Naturelle, Paris; and Patricia Vaurie before it was located. M.
Bonfils subsequently collected a series of topotypes and generously gave me
examples. |
56 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
LITERATURE CITED
Champion, G. C.
1911. Otiorhynchinae Alatae. In Biologia Centrali- Americana,
Coleoptera, Vol. 4, pt. 3, pp. 178-354.
Fall, “He €.
1907. Descriptions of new species. In Cockerell, T.D.A., and H.C.
Fall. The Coleoptera of New Mexico. Trans. Amer. Ent. Soc. 33:
145-272.
Fleutiaux, E., and A. Sallé.
1889. Liste des Coléoptéres de la Guadeloupe et descriptions d’especies
nouvelles. Ann. Soc. Ent. France 9: 351-484.
Howden, A. T.
1959. A Revision of the species of Pandeleteius Schénherr and
Pandeleteinus Champion of America North of Mexico. Proc. Calif.
Acad. Sci. (4) 29: 361-421.
Howden, A. T.
1963. A new species of Isodrusus, with notes on Isodrusus debilis
Sharp. Coleop. Bull. 17: 43-46.
Hustache, A.
1929. Curculionides de la Guadeloupe, pt. 1. In Faune des Colonies
Francaises, Tome III, fasc. 3, pp. 165-267.
Leng, C. W., and A. J. Mutchler.
1914, A preliminary list of the Coleoptera of the West Indies as re-
corded to January 1, 1914. Bull. Amer. Mus. Nat. Hist. 33 (30):
391-493.
Howden: West Indian Tanymechini D7
i,. 8. coccolobae feed-
Figs. 1-5. Scalaventer feeding damage on leaves.
ing on Coccoloba tenuifolia; 2, S. montanus feeding on Clethra occidentalis;
3. 8. cyrillae fee feeding on Cyrilla_ racemiflora; 4, S. litoreus feeding on Krugi -
dendron ferreum; 5, S. litoreus feeding on Haemotoxylum campechianum.
08 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
Figures 6-12. Scalaventer species.
Figs. 6, 8, 10, 11 lateral view: 6, cyrillae female; 8, coccolobae female
from Mahogany Vale; 10, montanus male; 11, gelinasus female.
Fig. 7, dorsal view, cyrillae allotype, female.
Fig. 9, 12, anterior view of head: 9, gelinasus female; 12, valkyrius
female.
60 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
Figs. 13-18. Abdomen of Scalaventer species: 13, coccolobae female
from Mahogany Vale; 14, litoreus female; 15, caymani holotype, female;
16, cyrillae male; 17, montanus allotype, female; 18, cubensis holotype,
male.
Howden: West Indian Tanymechini 61
Figs. 19-24. Scalaventer species. Figs. 19, 21, 23, lateral view of holo-
type: 19, jamaicensis male; 21, convexifrons male; 23, valkyrius female.
Figs. 20, 22, 24, abdomen of holotype, female: 20, remotus; 22, gelinasus;
24, valkyrius.
62 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
Figs. 25-31. Figs. 25-29, Paululusus species: 25, constanzae male,
dorsal view; 26, constanzae, anterior view of head; 27, calypso female, dor-
sal view; 28, hispaniolae female from San José de las Matas, anterior view of
head; 29, calypso female, lateral view. Figs. 30-31, Paradacrys species:
30, ensiformis allotype, female, from Castle Island, lateral view; 31, spatu-
latum allotype, female, anterior view of head.
Howden: West Indian Tanymechini 63
Figs. 32-37. Figs. 32, 33, Isodrusus guajavus: 32, lateral view; 33,
abdomen, female. Figs. 34, 35, Paradacrys spatulatum: 34, lateral view,
allotype, female; 35, abdomen, female, from Green Cay. Fig. 36, Pandeletei-
us testaceipes male from Guadeloupe. Fig. 37, Paradacrys ensiformis, ab-
domen of allotype, female.
64 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
Figures 38-45.
Figs. 38-41, Scalaventer species, lateral view: 38, litoreus female; 39,
remotus holotype, female; 40, caymani type, female; 41, cubensis holotype,
male.
Figs. 42-43, Scalaventer species, anterior view of head: 42, cubensis;
43, remotus.
Fig. 44, diagram of cross-section of abdomen through median line: a,
Scalaventer cyrillae; b, Scalaventer coccolobae; c, Pandeleteius nodifer.
Figs. 45, 46, diagram of vestiture of elytra of Paradacrys, a, dorsal
view, b, lateral view: 45, ensiformis; 46, spatulatum.
Figs. 47, 48, diagram of elytral setae in lateral view of Scalaventer: 47,
remotus; 48, litoreus.
66 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
Figures 49-60.
Figs. 49-50, foreleg of male of Scalaventer: 49, cyrillae; 50, coccolobae
from Mandeville.
Figs. 51-53, Paululusus hispaniolae: 51, lateral view allotype, female;
52, lateral view of elytra of female from San José de las Matas; 53, dorsal
view of holotype, male.
Fig. 54, foreleg of Paululusus calypso male.
Fig. 55, foreleg of Paradacrys spatulatum allotype, female.
Fig. 56, foreleg of Isodrusus guajavus male.
Figs. 57-58, lateral view of anterior portion of: 57, Pandeleteinus lucidil-
lus; 58, Scalaventer convexifrons.
Figs. 59-60, antenna of male Scalaventer: 59, coccolobae; 60, cyrillae.
4 Doral).
59
68 Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
Figures 61-76. Aedeagus, lateral view and dorsal view of apex.
Figs. 61-67, Scalaventer: 61, cyrillae; 62, coccolobae; 63, litoreus;
64, cubensis; 65, montanus; 66, jamaicensis; 67, convexifrons.
Figs. 68, 70, 72, 74, Paululusus: 68, calypso; 70, hispaniolae from
Port au Prince; 72, hispaniolae from San José de las Matas; 74, constanzae.
Fig. 69, Isodrusus guajavus.
Figs. 71, 73, Paradacrys: 71, ensiformis; 73, spatulatum.
Figs. 75, 76, Pandeleteius: 75, testaceipes; 76, nodifer.
70
Map 1.
Map 2.
Map 3.
Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
Distribution of Paululusus n.g., Paradacrys n.g. and Isodacrys
Sharp.
Distribution of Scalaventer n.g., Pandeleteinus Champion, and
Exmenetypus Voss.
Distribution of Isodrusus Sharp.
MAP 1
PAULULUSUS
PARADACRY S$
ISODACRYS
MAP 2
SCALAVENTER *
PANDELETEINUS @
EXMENETYPUS fe
MAP 3
ISODRUSUS DEBILIS
ISODRUSUS INSULANUS
ISODRUSUS GUAJAVUS
12
Contrib. Amer. Ent. Inst., vol. 5, no. 5, 1970
DOMINICAN REPUBLIC
MAP 4 HAITI
PAULULUSUS CONSTANZAE *
PAULULUSUS HISPANIOLAE SSS
PAULULUSUS CALYPSO Ie
Map 4. Distribution of the species of Paululusus.
Vs
INDEX 73
abdominal modifications, 4, 9, 29,
45, 53-54
Acacia, 52
Alchornea latifolia, 8
Bahama Islands, 5, 27, 43, 45, 50,
52, 54-55
Baja California, 5, 22, 54
biology: of Isodrusus guajavus, 52;
of Pandeleteius nodifer, 47; of
Scalaventer coccolobae, 13-14; of
Scalaventer cyrillae, 10; of
Scalaventer litoreus, 16-17; of
Scalaventer montanus, 19; of
Scalaventer remotus, 21
calypso, Paululusus, 32, 34-36
Cassia emarginata, 47
caymani, Scalaventer, 5, 6, 22-24,
26
Clethra alexandri, 10
Clethra occidentalis, 8, 10, 18-19
Coccoloba, 4, 13, 19; C. longifolia,
12; C, tenuifolia, 12, 21; -C.
troyana, 12; C. schwarzi, 12
coccolobae, Scalaventer, 4, 6, 9,
10-14, 17, 19, 25-26
Colombia, 47, 49
constanzae, Paululusus, 34, 36,
38-40
convexifrons, Scalaventer, 5, 6,
26-27
Cuba, 5, 22, 32, 53, 54-55
cubensis, Scalaventer, 5, 31-32
Cyrilla racemiflora, 4, 8-10, 13, 19
eyrillae, Scalaventer, 4, 6, 7-10, 13
19
debilis, Isodrusus, 50, 52. 54
Dominican Republic, 5, 37, 39
Ehretia tinifolia, 15-17
ensiformis, Paradacrys, 41, 43-45
Exmenetypus, Pandeleteius, 54
faunal list, 52-53
Florida, 5, 21-22, 47, 54
gelinasus, Scalaventer, 5, 27-29
Grand Cayman, 5, 23, 53, 54
Grenada, 49, 53, 54
Guadeloupe, 48-49, 53, 54
guajavus, Isodrusus, 30, 50-52, 54
Haematoxylum campechianum, 15-17
Haiti, 39, 31
Hispaniola, 29, 33, 53-55
hispaniolae, Paululusus, 33, 36-38
Honduras, 47
insulanus, Isodrusus, 50
Isodacrys, 2, 33, 40-41, 54
Isodrusus, 3, 4, 29, 40-41, 50-52,
53-55
Jamaica, 4-5, 8-10, 12-13, 15-17,
18, 19, 21, 20, 26, 30, 46, 47.
60, 92, 33-55
jamaicensis, Scalaventer, 5, 6,
24-26, 27, 32
key: West Indian genera, 3; species
of Isodrusus, 50; species of
Pandeleteius, 45; species of
Paradacrys, 41; species of
Paululusus, 33; species of
Scalaventer, 5
kirschi, Pandeleteius, 49-50
Krugiodendron ferreum, 15-17
Lantana camara, 47
litoreus, Scalaventer, 5, 14-17, 19,
30
montanus, Scalaventer, 5, 16, 17-19,
30
Nicaragua, 46-47
nodifer, Pandeleteius, 46-47, 53, 54
ocular vibrissae, 2, 31, 32, 41, 54
Pachnaeus, 3, 53
Pandeleteinus, 4, 27, 33, 41, 54
Pandeleteius, 2, 3, 9, 22, 45-50,
03-04 ee
Paradacrys, 2, 3, 4, 40-45, 53-55
Paululusus, 2, 3, 4, 29, 32-40,
03-55
Pithecolobium dulce, 47
Polydacrys, 3, 53
remotus, Scalaventer, 5, 19-21, 30
St. Vincent, 49, 53, 54 |
sex: distinguishing characteristics,
Scalaventer, 2, 3-32, 52-55
spatulatum, Paradacrys, 40, 41-43,
52
subtropicus, Scalaventer, 5, 6,
21-22, 24, 54
sublineatus, Pandeleteius, 50
Tanymecus, 2, 19, 53
teratology, 10, 13, 16
testaceipes, Pandeleteius, 45, 48-50,
03, 54
Vaccinium meridionale, 10
valkyrius, Scalaventer, 5, 6, 29-31
Venezuela, 49
a
SF
whe
Z
is
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A REVISION OF THE FAMILY SYRINGOPHILIDAE
(PROSTIGMATA: ACARINA) |
By
John B. Kethle y*
ABSTRACT
The family Syringophilidae is revised at a generic level anda key to the
two named and fifteen new genera is given.
The named genera and their type species are: Syringophilus Hellér, 1880,
S. bipectinatus Heller, 1880; Picobia Haller, 1878, P. heeri Haller, 1878.
New genera and their type species are: Syringophilopsis, Syringophilus elon-
gatus Ewing, 1911; Aulobia, Syringophilus dendroicae Clark, 1964; Aulonastus,
A. pipili, n. sp.; Chenophila, C. branta, n. sp.; Selenonycha, S. baltoda,
n. sp.; Creagonycha, C. lara, n. sp.; Niglarobia, N. ereuneti, n. sp.;
Syringophiloidus, Syringophilus minor Berlese, 1887; Philoxanthornea, P.
anoa, n. sp.; Peristerophila, Syringophilus columbae Hirst, 1920; Ascetomylla,
A. gallinula, n. sp.; Trypetoptila, IT. casmerodia, n. sp.; Stibarokris, S.
phalacrus, n. sp.; Syringonomus, S. bulwerius, n. sp.; Torotrogla, T. mima,
n. Sp.
The following named species of Syringophilus are transferred to new genera:
8. hylocichlae Clark, 1964, 8. turdi Fritsch, 1958, S. fringillae Fritsch, 1958,
and S. passerinae Clark, 1964, to Syringophilopsis; S. totani Oudemans, 1904,
to Creagonycha; S. helleri Oudemans, 1904, and S. trouessarti Oudemans,
1904, to Niglarobia; S. zenadoura Clark, 1964, to Peristerophila; S. dryobatis
Fritsch, 1958, and S. zumpti Lawrence, 1959, to Picobia; and 8S. villosus
(Hancock, 1895), to Torotrogla. The synonmy S. elongatus (Ewing, 1911) (Ss.
icteridae Clark, 1964) is ee oonutied: The following species were not seen and
they cannot be assigned to a genus on the basis of their descriptions: S. anthi
1 Supported in part by the Department of Entomology, University of Georgia,
Athens, National Science Foundation (GB-8606); and PHS Grant no. 5 TO1 AI
00216 to The Ohio State University.
2 The Acarology Laboratory, The Ohio State University, 1735 Neil Ave.,
Columbus, 43210.
2 Contrib. “Amer. Ent. inst. ;.vol. 5; no. 6, 1978
Pritech, 1956; Si bisetatus Fritech, 1956; 5. qlandaria Fritsch, 1958; §.. major
Berlese, 1878); 5. troglodyus Fritseh, E95e.
The family Heterochelidae Tragardh is transferred from the superfamily
Cheyletoidea to Anystoidea.
CONTENTS
ABOURAC Iacse 3 Ga Se NU ON ISE cc Seer taanes ae ge ieee a ae oath aa RP ae aE cen gig ALT aaa
ACKNOWLEDGMENTS. TREN a ae es er eae Gea in REE Re ied caecum Sapam ee
INTRODUCTION (nec , ; ARLE oS la tla Aa ee age he
COLLECTION AND PREPARATION OF MATERIAL . Bite tee uation eo ne eas co wane
ER OU pce Set caramel yg act gS gaa, ieee gk ae ag tile
Historical Review. seo. es gee Bi Ni mica age alge: ig gated Sigurt aaa
Systematic position of Sycncouhiidas 4 in ‘Chayletoidea.. eee aS ac amin *
PRA £8 Ge a gh tte ho Bee ee oa
Characters and descriptive methods, =: . . . ... ee
Gnathosoma , hs CC ante etd ae he RS gc es wk
Pee ag ass a Re Niklas rider we ee ae cares) Was ee Bemis. ee gree Ci tee dae he ec cas Sane a
LegS« «© ss ia eae Sioa hee yr cea CP ait “Roe see Remedy To
Key to the es of errinconMlidae. BRC eee Ue aia ee aC ig HOR ake oa hae ne
SS CI ee aaa i Se gO ee ce igs EE cart ae ececi b Sadho wt a ee ab a See ake wl - Se Y Senda: Ghee Se
Syreicopmiue pinectinatus Heller s,s no Bae ee ee a NO ee
Syringophiopsis. . . rag a Stee ee. Goat Ge Sar te ie Uae er ee eae
Syringophilopsis si jusees Swinal: Fe SS Sag ee al a as ae eee
Svringop nologies Mingia Writson.° 6 are eee he eS Fa eS
Svringommibonsia=yviocionia (Clary, . Sia coos wh a a ee eee
Pyringomaoseis passerine (Clark... i fo. sae Sw Wee Se EV aa
Syringe ilopeic Purdus “(Fritsch fo 4S ee a Oa ee
Te Oi Fe Re, ee Reig a ge ge ae VOB acts, Sle grt ge a ate
eer ay Varma eo ae Shige a ale ee ee ee een bee we ee
re a ee Sere 5) Saw Wea ee gece a a ag SR
BrepelOo 1a Caameroc a 6 ee a ea eee ee eee es Pe le a Ga
Torotrogla. . Pee wate ee oe Sg Se Mie meee. Le PU aR ee ce ears camrenres nes
Torotrogla villosa liancodh) big e Manes nti Se ae CRE aie cio 2S, Quen aa
ee i ee ae eo we ge wee el he oe a ee ee
Selenonycha. ... etretnge Or ute ge Ge” eh Sb ee athe ok a CO Cee Lee ee
Selenonycha Aaoda:. Rig ie ear aaa er gh oy ae ig ge eee ag gc omgetnag ga ak ee ge
RO ay ee el eg ete, ae i pees Be RR aR ok eg peg es See
Creagonycha lara. ... SREY AR gS ak age gta egg igh sag “age eg tg inera a
Creagonycha totana (Oudemans). Teo Se ae ee ea i Ne eC Ctr
ee a ee a gg eee Salhi es gS Bee eee
Orr eae em a a ee a we i ye aay a ya fe se
NMDWDOO A HP LP PW FF
Aulobia , oie setae We Wat han sary ltr Roe Goat a ling wag RQ oe
Aulobia dendreicns iCfarky .: hs cern ete RG meme be) oe Ne eee ae
Niglaropia, os, Pad ee ar nnrm ie a aha I keh a.) ae ued iia ce aa
Niglarobia apanaatl Grr ee ha ameter matte Pia curt sg OI gal ob agtallg dl ye“ Rigtaanep hal 3 a
Di eyo ie ear fF Ogerene ge) ee ae
Niglaronia Irouecemor.. (OUudemane) Ln a ce cece Sek ee we ee AF
Aulonastus e e e e e e e e e e e e e e @ e @ e e e e e e e e e e e e 47
PL Ue AEE Ph gr EE gee ea eee gran gs Wee iL Rees Ea ee ee
Syringopnifoidus o.oo. % std Nit aa pig tery ga tae lige agit: hike AY nh Taare
Syringophiloidus minor ‘@aniedei stag ane Nae lana gia eee hs. Clive ial ge hg eae
SePEG Ope te ie ee ae ari er i he ee ee
Kethley: Syringophilidae 3
PHISOMARUAOTIO Od io ee acini cael Mia aD ee ees RRS aE Weiew o ice, @e ie ae
Eee dL pe a OE a a ca ey le alg ae) Saltese uno est RR ee Bs igh ly he gia
Periaterophhlat-.n) 4. om Ue. shia Ley a CURL 1s Seas Bas eer wate 6 SS
Peristenophile..columba, Anaahat... godt are. 6 ee eR Er iow, BS
Peristerophila ,zenadoure Gland os ..iiges Sercdicisst otto seit,
SYTINGONOMUS 0) pean a ccetente e rehweee dl io cxosk ae aie Soar athe eee
Syringonomus- bu lwerius) so cleus myers 3h La oe oe Sei a te
Stibarokrig. a) wince tesnthlaaptanntil te Gdeo em ures UC en Seat 4s Soiree edie
Stibarokris, phelacrug.« ):+ 0-2 quien aiseoteh WORE ee ae eS RO. Gat Giese
PIOGHIG «. o.-6 - ee eh he ee we OW, at ei ie eal de oa Gave Bee a ed Eee
Picobia. heerk FB aer ot. ree becks Wy etuwelarceidet Le east, Beh soho whiete a44%. «04
Picobia drvobatic drrtisetiie: « sods Uleten cin eoekeleieae’ © Leneetsim 205
Picobia zumpti.. Lawrence) «dine likes sei cigdteouciat & een ae Letts. fear Go
TCR ee © oe kl Gen es Hee CAR: Cale ne le lee eae a Veer ean eae eee
Syringophilus antaharitses oo% 1a Set ar et ee ONS Or es ok a ke we
Byringophilus bigetatic- Feige po 50! a ieee ee ee 2 a) gees hale Sk A ee ca Gay a
Svringohpilus glandarii. Priteciik sai + catiecint. excaone iain ens: Sakhete STEP) tee oO
Swringophilus, major Berries gic p 4b Wh @ 1S a Pe BO RIOT Ogle ane. OSS
Syvringopnilvs trogiodviis Fyitson... wit al lec! deer we boas tes elie eee G elie
OE er ete kes ocmerbcitg diye. pe Ri Se sme i ie coped tee be | bere eke ee
BIONOMICS.AND HOST-PARASITE. RELATIONS sai wale boris VY Selle el arrestee 289
ete Te AR a a lobe cat eae Haine eek eel we ewes aes eis vel Beer Sts
Pilea etn Gs Leto cad. eth sehie obe. Aiea Bee op Ere btu areata. bs ya aie aol
Re AI a ate cas Wik. Ges shim lai ek «bly Sen Tee eatiee si ead isnahes Ate pies SE RA Le ei eee see
PAGAL is, Stine -cdie Si oie ar late. wea ables es 6d tit
ACKNOWLEDGMENTS
The writer wishes to express his sincere appreciation to Preston E. Hunter
and Warren T. Atyeo, (University of Georgia), for their guidance and encourage-
ment.
The following institutions made facilities available and/or arranged for the
loan of materials: British Museum Natural History (K. H. Hyatt); Laboratoire
d'Acarologie (M. Vachon, M. Naudo); Natal Museum (M. Lamoral); Department
of Health, Education, and Welfare Public Health Service, Rocky Mountain Lab-
oratory (C. E. Yunker); The South Australian Museum (D. Lee); United States
National Museum, Smithsonian Institution, Pacific Ocean Biological Survey
Program (A. B. Amerson); University of Georgia (P. E. Hunter); University of
Kansas (J. Knox Jones, Jr.); University of Nebraska (W. T. Atyeo, P. Peterson);
Zoologische Sammlung des Bayerischen Staates (E. Popp).
The author wishes to acknowledge deep appreciation to J. Gaud and C. D.
Radford who made personal collections available for study. The author is grate-
ful for the generous assistance of many unnamed collectors and contributors
of specimens.
4 Contrib. Amer. Ent. Inst., vol: 5, no. 6, 1970
INTRODUCTION
The cheyletoid family Syringophilidae Lavoipierre, 1953, includes a group
of twenty-four species of avian parasites. The genera of Syringophilidae are
now known from almost two hundred species of birds, representing sixteen
orders and forty-six families, from all the major geographic regions of the
world. This is a generic revision with new species described only as types of
new genera. Although only eleven new species are described, material repre-
senting over one hundred twenty-two species was examined; additional species
will be described in subsequent studies.
This study clarifies the relationships of the family Syringophilidae within
the superfamily Cheyletoidea and establishes a classification at the generic
level that can serve as a foundation for quill mite studies at the species level.
COLLECTION AND PREPARATION OF MATERIAL
Syringophilid specimens were acquired by field collections, loans, and
examination of forzen birds. Initial collections were made from preserved birds
in the Department of Entomology at the University of Georgia. All flight feath-
ers (primaries, secondaries, all coverts, alulars, and retrices) were removed
and examined. Heavy infestations of mites discolor the clamus and were easily
discovered by gross observation. Very low numbers, such as one or two indi-
viduals per quill, were detected under a dissection microscope using a strong
reflected light source.
Live mites and the infested quills were transferred to vials containing 70%
ethyl alcohol. Specimens preserved in alcohol for long periods required clear-
ing in lactophenol at 70°C for forty-eight hours. Similar treatment was re-
quired for rehydration of dead mites taken from frozen birds. Specimens were
mounted on microscope slides in PVA medium (Lipovosky, 1953; Beer, 1954).
This solution was found to be the most satisfactory for preparation of these
soft-bodied mites.
A phase-contrast microscope was used for the study of the slide mounted
specimens. Measurements were made with the aid of an ocular micrometer,
and drawings were prepared using a Wild microscope and drawing attachment.
TAXONOMY
HISTORICAL REVIEW
The family Syringophilidae was erected by Lavoipierre (1953) to include
those mites "armed with plates, leg I...unmodified, and the palp is well devel-
oped in some species." Prior to 1953, Syringophilus and Picobia were placed
in Myobiidae (Baker, 1949; Ewing, 1938). Lavoipierre transferred only Syringo-
philus to the new family, apparently being unaware of the existence of Picobia.
Actually, Picobia appeared in the literature from 1906 until the present only as
a name in checklists.
Dubinin (1957), unaware of Lavoipierre's paper, divided the complex of
Myobiidae into the families Harpyrhynchidae, Ophioptidae, Myobiidae, and
Syringophilidae. Dubinin characterized Syringophilidae (containing Syringo-
philus and Picobia) as: mites 3-3.5 times longer than wide, possessing
piercing chelicerae internal to a stylophore; with legs III and IV separate from
the anterior two pairs of legs, all legs being equally formed (Syringophilus) or
legs I and II more compressed than III and IV (Picobia), and living inside the
Kethley: Syringophilidae 5
quills of flight feathers or subcutaneous in the wing tissue of birds.
Although both Syringophilus Heller, 1880, and Picobia Haller, 1878, were
established many years ago, little work has been published concerning the na-
ture of these mites. Considerable confusion exists concerning the species, gen-
era and even the family. The only comprehensive treatment of Syringophilus and
Picobia was presented by Oudemans (1906). Various authors (Hancock, 1895;
Ewing, 1911; Lawrence, 1959; and others) described new species but gave in-
complete descriptions. Fritsch (1958) and later Clark (1964) described and
assigned additional species to Syringophilus. Unfortunately, only the holotypes
of four species and paratypes of six additional species are known to exist. All
of Fritsch's material was destroyed (in litt., H. J. Stammer) and Clark's holo-
types were lost in transit to the U. S. National Museum (in litt., E. W. Baker).
Prior to the present study, twenty-three species in the genus Syringophilus
and one species in the genus Picobia were assigned to the family Syringophilidae.
Inadequate literature reviews and insufficient generic definitions have led to the
incorrect placement of species by several authors (Fritsch, 1958; Lawrence, 1959;
and Clark, 1964). Critical examination of the twenty-three named species of
Syringophilus clearly indicated that the genus has become a confusing conglom-
erate of diverse forms. Excepting several works describing single species, the
majority of the literature references are checklist citations.
SYSTEMATIC POSITION OF SYRINGOPHILIDAE IN CHEYLETOIDEA
The superfamily Cheyletoidea is characterized as follows: cheliceral bases
completely fused with subcapitulum and forms a stylophore lacking a median di-
vision; movable chela straight and styletlike, or long and whiplike; peritreme
internal to the stylophore and usually M-shaped; a single median dorsal opening
present on the stylophore; adults lack pseudostigmata and genital discs; usually
with a palpal thumb-claw complex; free living or parasitic on vertebrates. The
families which lack the thumb-claw complex are more specialized morpholog-
ically and biologically. Among the parasitic species, the genital opening of the
male has migrated from a ventral position to a terminal and finally true dorsal
locus. Camin etal. (1967), noted that among prostigmatid mites, the dorsal
location of the male genital opening is restricted to the Cheyletoidea.
The following families are presently placed in Cheyletoidea:
Cheyletidae Leach 1814 free living, predaceous
Harpyrhynchidae Dubinin 1957 bird skin
Syringophilidae Lavoipierre 1953 bird quills
Myobiidae Megnin 1877 mammal skin
Psorergatidae Dubinin 1955 mammal skin
Demodicidae Nicolet 1855 mammal skin
Ophioptidae Southcott 1956 snakes
Chloracaricae Camin et al. 1967 turtle cloaca
Heterochelidae Tragardh 1950 beetles (Passalidae)
Baker et al. (1958) incorrectly credits Dubinin with the authorship of Syringo-
philidae and Ophioptidae. Heterochelidae Tragardh 1950, should be removed to
the Anystoidea (Cunliffe, 1955) due to the presence of a suture on the stylophore
and a pair of propodosomal pseudostigmata.
The high degree of specialization of the parasitic forms within the Cheyle-
toidea makes interfamilial relations difficult to ascertain. Modifications for para-
sitic life are so great as to obscure the relations of structures and/or setae among
6 Contrib. Ameratfntsiinsty, vol. 5, 70. 6, 1970
certain groups in this superfamily. The legs and idiosomal setation have been
reduced in Chloracaridae, Ophioptidae, Demodicidae, and Psorergatidae. My-
obiid mites have conspicuous elaborations of legs I. The remaining families -
Cheyletidae, Harpyrhynchidae, and Syringophilidae - are closely related. The
presence of segmented legs III and IV and ambulacra in species of Harpypalpus
Dubinin 1957, indicates an intermediate degree of specialization between cheyle-
tids and species of Harpyrhynchus Megnin 1878. Comparative chaetotaxy reveals
that the syringophilids are close to the cheyletids that are hibitually found in-
side feather shafts.
DIAGNOSIS OF SYRINGOPHILIDAE
Syringophilidae Lavoipierre 1953
The family Syringophilidae is characterized within the Cheyletoidea as
follows: elongate to oval, soft-bodied mites with long needlelike chelicerae
and complete stylophore; palpus four segmented lacking a well developed thumb-
claw complex; peritreme well developed; legs I-II distant from legs III-IV; am-
bulacrum composed of two claws separated by a bipectinate empodium; male
genito-anal opening dorsal; living in quills of flight feathers of birds or sub-
cutaneous in the epidermis of the wing.
CHARACTERS AND DESCRIPTIVE METHODS
Listed below are explanations of terms and relative measurements employed
as diagnostic or descriptive characters for the mite genera involved in the study.
The explanations parallel the sequence used in the formal descriptions.
Gnathosoma - characters 1-6 |
The gnathosoma consists of the stylophore, movable digits, hypostome, and
paired palpi. The movable digits of the chelicerae and the peritremes are inter-
nal to the stylophore (Fig. 1).
1. Hypostomal apex: Highly ornate (Figs. 10 C, D; 12 C, D), hypostome pro-
duced anteriorly into hyaline expansions and protuberances; slightly ornate
(Fig. 23 C, D), hyaline protuberances present, but inconspicouos; smooth (Fig.
22 C, D), hyaline protuberances absent.
2. Lateral hypostomal teeth: Present - lateral sclerotization of the hypostomal
lips protruding in the region of the adoral setae (Fig. 36 C, D): absent - scler-
otization of lateral hypostomal lips lacking, although an apodeme internal to
the surface of the adoral setae may be present.
3. Movable chela: Dentate - small, often minute teeth present on the distal tip
of the digit; edentate - lacking teeth.
4. Peritremes: Shape of the peritreme - "U" shaped - recurrent, or longitudinal
branches produced posteriorly, displacing the lateral branches (Fig. 8 C); "M"
shaped-recurrent branches not produced posteriorly (Figs. 14 C, 16 C). The
maximum and minimum number of chambers in each of the lateral and longitud-
inal branches is given for each of the species studied.
5. Stylophore: Shape of posterior portion of the stylophore: rounded - tapered
to a gentle curve (Figs. 18 A, 20 A, 27 A); constricted - with a constriction and
further elongation (Fig. 36 A). Relation to propodosomal plate: subcutaneous -
extending posteriorly under the propodosomal plate (Fig. 22 A); independant -
not extending posteriorly under the propodosomal plate (Fig. 1).
Kethley: Syringophilidae 7
Psc St, a
XN
\
. ve
\
SCi
Fig. 1. Lateral aspect of Aulobia dendroicus (Clark): Ch, movable
digit of chelicera; LL, lateral hypostomal lips; P, acetabulum of
palpus; Pl, longitudinal branch of peritreme; Psc, propodosomal
plate; Pt, transverse branch of peritreme; Sta, apodemal portion
of stylophore; Ste, exposed portion of stylophore; I, acetabulum
of leg I; II, acetabulum of leg II. aol, ao2 - adoral setae; vi, ve,
sci - propodosomal setae.
6. Condition of palpal apex: Rounded - distal margin of palpal tibiotarsus
rounded with omega ventarad of K', K" (Fig. 2 A); truncate - distal margin of
palpal tibiotarsus truncate with omega in the same plane as K', K" (Fig. 2 B).
Idiosoma - characters 7-17
A combination of signatures for the idiosomal setae has been used for the
dorsal propodosomal setae; Baker, 1965, for the dorsal hysterosomal setae; and
Grandjean, 1944 for the ventral idiosomal setae. The positions and correspond-
ing designations for the setae of the body exclusive of the legs are found in
Fig. 3A, Band Fig. 4A, B.
7. Setal types of idiosoma and legs (except a', a" of tarsi I-IV): smooth - seti-
form and hairlike (Fig. 27 C); annulate - circular or spiral sculpturing on the seta
(Fig. 34 C); knobbed--regularly arranged rounded raised portions on the surface
of the seta (Fig. 36 E); normal base--diameter of the seta at the base as large
or greater than the diameter slightly distad (Fig. 27 C); constricted base--diam-
eter of the seta at the setal base less than the diameter slightly distad
(Fig. 36 E).
8 Contrib. Amer. Ent: Inst.; vol. 5) 70. 6, 1970
Fig. 2. Dorsal aspect of left palpus: A, Distal margin of tibiotarsus
rounded, Syringophiloidus sp.; B, Distal margin of tibiotarsus trun-
cate, Picobia zumpti (Lawrence); omega-solenidion; K', K"-eupathids.
8. Condition of propodosomal plate: Divided--separated into right and left por-
tions; fragmented--disjunct sclerotization around the bases of the setae; not
divided: rectangular--margins relatively straight; variable--conditions of mar-
gins as scalloped, cleft, or concave.
9. Hysterosomal plate: Present--sclerotized in the region of setae 12, d3, 13;
absent--sclerotizion absent in the region of setae 12, d3, 13. If present, the
hysterosomal plate may fuse with the pygidial plate, see #12 below.
10. Setae in region of propodosomal plate: Six pairs of setae (vi, ve, sci, sce,
di, 11) or five pairs (vi absent) are associated with the propodosomal plate.
Configurations of the setae are translated into numerical patterns by counting
the vertical and scapular setae in a longitudinal direction, and dl, llina
transverse direction as: 3-1-2, Fig. 5 A; 2-2-2, Fig..5 GC; 2-1-1-2,.Fig..5 D;
3-2-1, Fig. 5 E. The patterns differ in the male and are not employed asa
character.
Kethley: Syringophilidae 9
oXe\ AO;
O O
dq
l4 5 9 G2. os
° ° re)
Mi |
° Ig
Fig. 3. Setal signatures of body. dorsum: female, A; male, B. aol -
adoral; al, a2 - anal series; dl, d3, d4, dS - dorsal series; gl, g2-
genital series; 11 to 15 - lateral series; sci - scapular internal; sce -
scapular external; vi - vertical internal; ve - vertical external.
10 Contrib. Amer. Ent. Inst., vol. 5, no. 6, 1970
i F
Ag
: 1
2 rs
SJ. oO
Ki) (es
Fig. 4. Setal signatures of body, venter: female, A; male, B. ao2 -
adoral; la, lb, le - coxa I; 2b ~ coxa Il; 3a, 3b, 3c - coxa III; 4b,
4c - coxa IV; gl, g2 - genital series; pgl, pg2, pg3 - paragenital
series; pmc - subcapitular; scl to sc4 - trochanters I to IV.
Kethley: Syringophilidae 11
2-\-I-2
D
Fig. 5. Diagrammatic representation of configurations of setae
associated with propodosomal region. See text, p.8, for
explanation.
tz Contrib).AmercbntsInst. vole Spanos: 6, 1970
11. Setae in region of hysterosomal plate (12,d3, 13): Relative lengths--leng--
equal to or greater than dl; short--conspicuously shorter than dl. Arrangement--
relative positions of the setae to each other as stated in the descriptions.
12. Relative lengths of terminal setae (a4, 14, d5, 15): Female--Long--equal to
or greater than dl; short--conspicuously shorter than dl; male--d5, 15 absent,
d4, 14 as infemale. Any combination of these setae may be borne or a weakly
sclerotized region termed the pygidial plate.
13. Genital and anal setae: one or two pairs each of genital and anal setae
present in either the male or female.
14. Paragenital setae: Female--typically with three pairs; male typically with
two or three pairs; additional pairs (1-6), termed supernumerary, may be present.
The stated number of setae includes the entire complement, i.e., the basic
plus supernumerary members. :
15. Condition of coxal apodemes I and II: Parallel--mesal coxal apodemes of
legs I (MCA1) parallel to each other (Fig. 6 A); divergent--mesal coxal apo-
demes of legs I (MCA1) curve laterally along posterior 1/2 of length (Fig. 6B,D);
convergent--mesal coxal apodemes of legs I (MCA1) curve mesally toward each
other along posterior 1/2 of length (Fig. 6 E): degree of curvature--strongly di-
vergent (Fig. 6 D), or weakly divergent (Fig. 6 B, C); fused--union of mesal
coxal apodeme of leg I (MCA1) and mesal coxal apodeme of leg II (MCA2) prom-
inent (Fig. 6 B, C, D): point of fusion--MCAI1 fused to MCA2 at posterior 1/2
of MCA2 (Fig. 6 B), MCA] fused to MCA2 at anterior 1/2 of MCA2 (Fig. 6 C);
free--union of mesal coxal apodeme of leg I (MCA1), to mesal coxal apodeme of
leg II (MCA2), lacking or imperceptable (Fig. 6 A, E).
Fig. 6. Region of coxae I-II, female: A, Syringophiloidus minor; B,
Syringophilus bipectinatus; C, Syringophilopsis elongatus; D, Sele-
nonycha baltoda; E, Picobia sp.-MCA1l-mesal coxal apodeme leg I;
MCA2 mesal coxal apodeme leg II.
Kethley: Syringophilidae 13
16. Extent of sclerotization of coxae II] and IV: Weakly sclerotized--margin of
coxa merges with integument; strongly sclerotized--margin of coxa distinct.
17. Direction of cuticular striation: Longitudinal--in the direction of the meson;
transverse--in a lateral direction; completely transverse--at least one striation
connecting left pleuron to right Sap # incompletely transverse--striations
curved toward meson.
Legs - characters 18-21
Each leg consists of coxa, trochanter, femur, genu, tibia, tarsus, and pre-
tarsus. Although the shape of the segments may vary, segments are never sub-
divided or lost. The chaetotaxy of the legs (Fig. 7, Table I), is stable fora
given genus and affords a very useful character. Grandjean's (1944), chaeto-
taxal signatures for stigmaeids has been applied in the current study with only
minor modifications.
18. Thickness of legs: Relative thickness of legs as contrasted with each other
from the femur to pretarsus.
19. Leg chaetotaxy: Full complement of setae--scl, sc2, dFII, dFIII, dFIV,
dGII, dTIII, driv, vs'I, vs"II present; reduced--any or all of the foregoing setae
absent; supernumerary--vsIII present.
20. Condition of a', a" (fan setae): multiserrate--distal tip expanded, bearing
four or more regularly spaced serations (Fig. 32 C); rod-like--distal tip not ex-
panded, but rounded, and of uniform diameter throughout (Fig. 37 D); irregularly
serrate--distal tip bearing three or fewer irregularly spaced serrations (Fig. 37E).
21. Condition of claws: paraxial and antaxial members sub-equal; paraxial and
antaxial members dissimilar (Fig. 37 C); recurved--distal portion of claw curv-
ed back toward basal portion, basal portion enlarged (Fig. 20 C); broadly curv-
ed--"C" shaped, distal portion not reflected upon basal portion, basal portion
not enlarged (Fig. 18 C); with basal angle--basal portion of claw forming a
right angle to curvature of the main body of claw (Fig. 25 C); relative length of
Cclaws--length of claws as a fraction of the length of the empodium.
In the generic descriptions, the characteristic number, i.e., (1)., (2).,
(3)., etc. is underlined for distinctive characters for that genus.
14 Contrib. Amer. Entadnst.- voles'5;-no.-6, .1970
TABLE I
Leg chaetotaxy of
Syringophilus bipectinatus Heller adults
(Adapted from Grandjean, 1944)
Number of
Segment setae Setal signature
Coxae I-IV ce SR, Ly tg ba Ao Shin MO. Gos eke
Trochanters
I-IV beste lo Bele Bee. Sco, Bue.
Femur I 2 OF, vF.
II 2 iy Vis
III l OF.
IV 1 dF.
Genu 3 dG. Hay sigma,
II 2 daG, 1G
III ] al
IV l 1G"
1itte 4 5 Cl Weeks mo” yo ed.
I 4 ar, 0, 1.
III 3 GL Wi gelt
IV 3 Cly Mieete 3
Tarsus
I Ae u', us vs', ft, eae eee P', Pos a’, ae omeg e
iu 8 u', u", vs’, te’, te", a’, a”, omega.
III 6 a’, w, fe", te", 2's as
IV 6 we. ee cee ree
‘denotes paraxial position.
" denotes antaxial position,
Solenidia included in total setal count per segment.
Complete uppercase lettering denotes eupathids.
Kethley: Syringophilidae Ld
Pig. 7. Schematic dorsal aspects of legs I-IV. Dorsal tactile setae
represented by open circles, dorsal solenidia by incomplete circles,
ventral tactile setae py solid black circles.
16
LO,
els
ie;
Contrib. Amer, Ent. Insts; vol. 5; no. 6, 1970
KEY TO THE GENERA OF SYRINGOPHILIDAE BASED ON ADULT FEMALES
Palpal tibiotarsus truncate on distal margin; setae a' anda" of tarsi not
multiserrate; legs I and II dissimilar to legs IIIT and IV ........cc.ceeeees
sae Se eso ee meee eb ele ese es ee ee eee ences Picobla Haller, ». 64.
Palpal tibiotarsus rounded on distal margin; setae a' and a" of tarsi with at
least four serrations; legs I and II similar to legs III and IV............2.
Dorsal sétee Of femur ll (AFIT) abeGent .. occ cece ccc csc cs ccccsb Sees Oe
Dorsal setae Of omur TT (GFT) resent §. wo. ccc cc cca sce accecess 4
Setae vs'II absent; vi absent; Columbiformes .....cccrcccsccccsccsesecs
idee 6666s be cos oo. sl Perisieronnile, new genus, Pp. 53.
Setae vs'II Ree ene Vi present Charadritformes .. 3 6 secs ccs bole se le
Se Gs Ree a 6 See a es bis be Ss Oe ees we .. Ascetomylla, Adw aerue. x ‘4.
Dereal Setes Of genu 1] (GID absent coc oink. doesn cee ete cic an ee Oe
Dorsal setae of genu II (dGII) present .....c00000 sia tsckaeeealas er ns ac
Setae vs'I absent; Pelecaniformes and Charadriiformes ....... oes ae es
Pe le ee Oe ae ee G Spe eee See 6 Philoxanthornea, new sonud: p. 30.
Setae vs'I present; Passeriformes ...Syringophiloidus, new genus, p. 48.
At. least four pairs of paragenital setae present .... cc cervcvvcscsccee se
No more than three pairs of paragenital setae present...... wie WHE, Boarmanten Oe
Setae vs'III present; at least seven pairs of paragenital setae present;
large mites; Ciconiformes............... Irypetoptila, new genus, p. 25.
Setae vs'III absent; at most five pairs of paragenital setae present; moder-
ate sized mites; Passeriformes............ Lorotrogla, new genus, p. 28.
One pair of anal or one pair of genital setae absent ........ccecerveee Ge
Two pairs of anal setae present and two pairs of genital setae present. 10.
One pair of genital setae present; two pairs of anal setae present; vs'II ab-
sent; vi present; Procellariformes.......Syringonomus, new genus, p. 56.
Two pairs of genital setae present; one pair of anal setae present; vs'Il
present; vi absent; Passeriformes......... Aulonastus, new genus, p. 47.
Ses ke ig wb ie ob beh ace oo wie boosie @eeee#ee# pe ee aed | ae
Setae vc 1 Present 0. ben ek se ewe ee Se uc pers eae bose ce aes
Lateral hypostomal teeth present; propodosomal setae 2-2-2; Coraciiformes
6 4s Wile Bias up eteieiceinsraloasls aie cohen edwas foe wale « DELDONOKELS.« NOW+GENUS +>: Pee 62s
Lateral hypostomal teeth absent; SID BOABE Seal SOtae 2-4 also s ive seme os 8
eee UU ee ee bee ee ieee os sey eka sae N@larObia, New genus, p. 43.
Beta 12, do, 13, do, Jeng 4 c.ees cee uue Caisse ees ese ERD a aap cbos
Becae 12.03, 13, do, Short) ANSerilOrmes. . 2b se oe a ec oe sc he cee see a ae
ree ee Ppas es be Noe a RUA oe cee bes oes CeNODnia, New. genus, Dp. .o7.
Kethley: Syringophilidae 7
13; Mesal coxal apodemes of legs I (MCA1) parallel; Passeriformes ..........
aerate ere irs es FE i weocvas Bulobi new genus, p. 4l.
Mesal coxal apodomes of legs I (M CAl1) divergent, not parallel.......14,
14. Mesal coxal apodemes of legs I (MCA1) dissimilar in size and shape of
mesal coxal apodeme of legs II (MCA2); claws at most 1/3 length of
embpoOditin=. fob 78s a a Fe oa VER COE Ea be Os a Os a
Mesal coxal apodemes of legs I (MCA1) similar in size and shape to mesal
coxal apodeme of legs II (MCA2); claws at least 1/2 length of empodium 16.
15. Hypostomal apex unornamented; mesal coxal apodeme I (MCAI1) fused to
mesal coxal apodeme II (MCA2) at posterior base of MCA2; d2 usually
closer to 12 than to d3; Galliformes..........Syringophilus Heller, p.17.
Hypostomal apex ornamented; mesal coxal apodeme I (MCA1) fused to mesal
coxal apodeme II (MCA2) at anterior 1/2 length of MCA2; d2 usually closer
to d3 than to 12, but may be equidistant between 12 and d3; Passeriformes.
CC este ree ese s towews see cb es oe ees sD VIInGOD MlOpsis, new genus, Pp. ES:
iG. Claws’ recurved; dso short; Characriiiormes. s 5s sss es a a eee a ee ees
a eee es ee eas bs es kee e «ee ee Creagonycha, new genus, p. 34.
Claws’ broadly opew?.do long; ‘Charadrirormes 6.6. 6 rs 5 ccs soe sae vce eles
egw aha erent Cee e ar ee Wecteesesscone ss sOelenonvena, Tew cenus, p.. 31.
Genus Syringophilus Heller
Syringophilus Heller, 1880, Die Schmarotzer, 186 - Oudemans, 1906, Mem.
Soc. zool. Paris, 19:43-46. - Hering, 1934, L'Amateur do Papillons, Paris,
7, 154 (Hymenoptera: Insecta, not Acari). - Dubinin, 1957, Mag. Parasitol.
Inst. Zool. Akad. Sc. USSR., 17:71-136. - Fritsch, 1958, Zool. Jahrbucher
86:227-229.
Within the group of genera having MCA] weakly divergent and fused to MCA2
(Syringophilus, Syringophilopsis, Ascetomylla, and Trypetoptila) Syringophilus
is distinguished by the configuration of the lateral branches of the peritreme,
and the location of setae a3 closer to 12 than to13. This genus appears most
similar to Syringophilopsis. These large mites (920u- 1100y) are found in the
flight feathers of galliform birds of the families Phasianidae and Numididae.
Type species: Syringophilus bipectinatus Heller, 1880, Die Schmarotzer,
p. 186, (by original designation); ex. Gallus gallus (L.); Phasianidae, Galliformes.
FEMALE, (1). Hypostomal apex smooth. (2). Lateral hypostomal teeth
absent. (3). Chelicerae dentate, 1-2 small teeth. (4). Lateral branches of
peritreme displaced into U shape, (Fig. 8 C), with 2-3 chambers; each longi-
tudinal branch with 8-10 chambers. (5). Posterior margin of stylophore pointed,
extending beneath propodosomal plate. (6). Palpal tibiotarsus rounded on dis-
tal margin. (7). All setae smooth. (8). Propodosomal plate weakly sclerotized;
lateral margins parallel; anterior and posterior margins concave. (9). Hystero-
somal plate absent. (10). Setal pattern of propodosomal region with six pairs
of setae arranged 3-1-2. (11). Setae 12, d3, 13 long; d3 closer to 12 than to 13.
(12). Setae d4, 14 long; d5, 15 short. (13), eating es lee with two pairs of
setae; anal series with two pairs of setae. (Way: Paragenital series with three
pairs of setae. (15). MCAl weakly divergent fused to MCA2 at posterior base of
MCA2. (16). Coxae III and IV subequal, crescent shaped, weakly sclerotized,
margins indistinct. (17). Cuticular striations as in Fig. 14. A, B. (18). Legs
18 Contribs Amer: Enté-dInsts, vols: 5,.no. 6, 21970
I and II thicker than III and IV. (19). Legs with full complement of setae. (20).
Setae a' anda" multiserrate, 6-15 tines. (21). Antaxial and paraxial members
of claw pair subequal; claws 1/3 to 1/4 length of empodium. (22). Order of
hosts: Galliformes. (23). Types of feathers inhabit: unknown.
MALE. As in female except: (3). Chelicerae edentate. (10). Setal pattern
of propodosomal region 3-1-2, all setae on plate. (11). Setae d3, 13 short.
(12). Setae d4 short. (14). Paragenital series with two pairs of setae; pgl
absent, pg3 terminal.
Remarks. In addition to the type species, material representing one un-
named species was studied in an evaluation of the generic characters.
Syringophilus bipectinatus Heller
(Figs. 8, 9)
Only the pertinent citations of this species are given.
Syringophilus bipectinatus Heller, 1880, Die Schmarotzer, 186. - Oudemans,
1906, Mem. Soc. zool. Paris, 19:47-57. - Baker and Wharton, 1952, An
introduction to acarology, 229-231. - Lavoipierre, 1953, Trans. Roy. Soc.
Trop. Med. Hyg. 47:7. - Dubinin, 1957, Mag. Parasitol. Inst. Zool. Akad.
Sc. USSR., 17:71-136. - Fritsch, 1958, Zool. Jahrbucher 86:229-230. -
Clark, 1964, Acarologia 6:77.
Although all type material is lost, 16 specimens on loan from the British
Museum (Natural History), adjudged to be conspecific were examined by the
author. Described originally from Gallus gallus (L.), (Galliformes: Phasiani-
dae), in Europe, this species has been collected from the same host in Ndumu,
Zululand. and from Kansas and Colorado in the United States. Numerous records
of this mite appear in the literature from different hosts, but only the above
listed reports have been confirmed by the author.
Syringophilopsis, new genus
Within the previously mentioned group of genera having MCA] fused to MCA2,
species of Syringophilopsis may be distinguished by the shape of the lateral
branches of the peritreme, the location of d3 closer to 13 than 12, and the pres-
ence of the full complement of leg setae. This genus appears most similar to
Syringophilus. These large mites (950u - 1360) are found in the primaries of
passeriform birds of the following families: Aludidae, Fringillidae, Hirundinidae,
Icteridae, Parulidae, Pycnonotidae, Turdidae, Tyrannidae, and Vireonidae. The
name Syringophilopsis (masculine singular) means having the appearance of
syringophilus, and refers to the similarity of the two genera.
Type species: Syringophilus elongatus Ewing, 1911. Psyche, Boston 18, 39.
FEMALE. (1). Hypostomal apex highly ornamented; 2-3 paired median pro-
tuberances. (2). Lateral hypostomal teeth absent. (3). Chelicerae dentate,
1-3 teeth. (4). Peritreme M-shaped; longitudinal branches each with 14-20
chanbers; lateral branches each with 2-6 chambers. (5). Stylophore rounded
posteriorly; not extending beneath propodosomal plate. (6). Palpal tibiotarsus
rounded on distal margin. (7). All setae smooth. (8). Propodosomal plate
divided or deeply cleft on anterior margin. (9). Hysterosomal plate variable.
(10). Setal pattern of propodosomal region with six pairs of setae arranged
3-1-2; 11 not on plate, variable position. agli Setae 12, d3, 13 long; d3 closer
to 13 than 12, or equidistant between 12 and13. (12). Setae d4, d5 long; 14, 15
Kethly: Syringophilidae
lOOp
Fig. 8. Syringophilus bipectinatus Heller. Female: A, dorsal aspect;
B, ventral aspect; C, peritreme.
ee
- 320
Contrib, Amer. Ent: inst. ; vol. 5, no. .6,.1970
¢ s
i
\
\
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oe
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Cee
Pig. 9. Syringophilus bipectinatus Heller, Male: A, dorsal aspect;
B, ventral aspect.
Kethley: Syringophilidae 21
variable. (13). Genital series with 2 pairs of setae; anal series with two pairs
of setae. (14). Paragenital series with three pairs of setae. (15). MCA] diver-
gent, fused to MCA2 at anterior 1/2 length of MCA2. (16). Coxae III and IV moder-
ately sclerotized, rectangular in shape. (17). Cuticular striations as in Fig.
10 A, B. (18). Legs I-IV subequal in thickness. (19). Legs with full comple-
ment of setae. (20). Setae a' anda" multiserrate, 12-20 tines. (21). Antaxial
and paraxial members of claw pair subequal; claws 1/3 length of empodium.
(22). Order of hosts: Passeriformes. (23). Types of feathers inhabit: primaries.
MALE. As in female except: (1). Hypostome without hyaline lips. (3).
Chelicerae edentate. (8). Propodosomal plate very weakly sclerotized, margins
indistinct. (9). Hysterosomal plate absent. (10). Setal pattern of propodo-
somal region obscured, difficult to characterize. (11). Setae 13, d3 short. (12).
Setae 14 long; d4, d5, 15 short. (17). Cuticular striation patterns as in Fig. 11
Ay By
The genus Syringophilopsis includes five named and sixteen new species.
The named species to be transferred to this genus are:
Syringophilopsis elongatus (Ewing)
new combination
Figs. 10,1)
Syringophilus elongatus Ewing, 1911, Psyche, 18, p. 39. Syringophilus icteridae
Clark ,.19604, Acarologia 6: 89, Figs. 30,31.
Although all type material is lost, 50 specimens on loan from the Rocky
Mountain Laboratory, from the type host, adjudged conspecific were studied.
Described originally from Agelaius phoeniceus (L.) (Passeriformes: Icteridae)
this species has been collected in Maryland and Georgia, U.S.A.
Syringophilopsis fringilla (Fritsch)
new combination
Syringophilus fringillae Fritsch, 1958, Zool. Jahrbucher 86 (3):230-233, Figs.
2,2.
Although all type material is lost, 3 specimens on loan from C. D. Radford
and J. Gaud from the type host were adjudged conspecific. Described originally
from Fringilla coelebs (L.) (Passeriformes: Fringillidae) in Germany, this species
has been collected from the same host in England and from Carduelis carduelis
(L.) (Passeriformes: Fringillidae) in Morocco.
Syringophilopsis hylocichla (Clark)
new combination
Syringophilus hylocichlae Clark, 1964, Acarologia 6: 87, Figs. 28, 29.
Although the holotype is lost, 2 paratypes on loan from the Rocky Mountain
Laboratory were examined. Described orginally from Hylocichla fuscescens
(Stephans) (Passeriformes: Turdidae) from the United States, this species has
not been recollected.
22 Contrib. Amer. Ent. Insts, vol..5y)-10..6,..1970
lOOu
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WS)
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Wei
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Sa
WO
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Fig. 10. Syringophilopsis elongatus (Ewing). Female: A, dorsal
aspect; B, ventral aspect; C, hypostomal lips, dorsal aspect; D,
hypostomal lips, ventral aspect.
Kethley: Syringophilidae 23
Fig. 11. Syringophilopsis elongatus (Ewing). Male: A, dorsal
aspect; B, ventral aspect.
Syringophilopsis passerina (Clark)
new combination
Syringophilus passerinae Clark, 1964, Acarologia 6: 91 Figs. 26, 27.
Although the holotype is lost, 1 paratype on loan from the Rocky Mountain
Laboratory was examined. Described originally from Passerina cyanea (L.) from
Maryland, this species has been collected from the same host in Florida, U.S.A.
24 Contrib. Amer. Ent. Inst., vol.-5;,no.. 6, 1970
Syringophilopsis turdus (Fritsch)
new combination
Syringophilus turdi Fritsch, 1958, Zool. Jahrbucher 86(3):233-235, Fig. 4.
All type material of this species is destroyed, and representative specimens
were not avilable for study. The type host of this species is Turdus pilaris (L.)
(Passeriformes: Turdidae) in Germany.
Ascetomylla, new genus
Among the previously mentioned genera having MCA] divergent and fused to
MCA2, Ascetomylla is distinguished by the configuration of the hypostomal lips
and the loss of leg setae dill, dFIII, and dFIV. Based upon the epimerites, this
new genus appears closely y related to Syringophilopsis. These large to very
large mites (1200p - 1340) are found in the flight feathers of gruiform birds of
the family Rallidae, and coraciiform birds of the family Bucerotidae. The name
Ascetomylla (feminine singular) means ornamented lip and refers to the highly
ornate condition of the hypostomal lips of the adult female.
Type species: Ascetomylla gallinula new species, ex. Gallinula chloropus
(L.); Rallidae, Gruiformes. ;
FEMALE. (1). Hypostomal apex highly ornamented, two fingerlike and two
rounded, paired protuberances present (Fig. 12 C, D). (2). Lateral hypostomal
teeth absent. (3). Chelicerae with 2-3 teeth. (4). Peritreme M-shaped; lat-
eral branches each with 3-5 chambers; longitudinal branches each with 16-24
chambers. (5). Stylophore rounded posteriorly, to weakly constricted; not
extending beneath propodosomal plate. (6). Palpal tibiotarsus rounded on dis-
tal margin. (7). All setae smooth. (8). Propodosomal plate divided; lateral
margins concave and scalloped. (9). Hysterosomal plate variable. (10). Setal
pattern of propodosomal region with six pairs of setae arranged 3-1-2. (11).
Setae 12, d3, 13 long; d3 closer to 13 thanto12. (12). Setae d4, 14, 15 long;
subequal; 5. short; 14, 15 borne on common tubercle. (13). Genital series with
2 pairs of setae; anal series with two pairs of setae. (14). Paragenital series
with three pairs of setae. (15). MCA1 divergent, fused to MCA2 at anterior 1/3 to
1/2 length of Ep2. (16). Coxae III and IV weakly sclerotized, margins indistinct.
(17). Cuticular striations as in Fig. 12 A, B; transverse between coxae III and
IV. (18). Legs I slightly thicker than II; legs II subequal in thickness to III and
IV. (19). Leg setae dFII, dFIII and dFIV absent. (20). Setae a' anda" multi-
serrate, 16-24 tines. (21). Antaxial and paraxial members of claw pair sub-
equal; claws 1/2 length of empodium. (22). Orders of hosts: Gruiformes and
Coraciiformes. (23). Types of feathers inhabit: unknown.
MALE. As in female except: (1). Hypostomal lips lacking. (8). Sclero-
tization present in region of setal bases vi, ve only. (9). Hysterosomal plate
absent. (10). Setal pattern of propodosomal region obscure. (11). Setae 12,
‘d3; 13 short. (12). Seta 14 long; d4, d5, 15 short: (17)... Cuticular. striations
as in Fig. 13 A, B. (21). Claws 1/3 length of empodium.
Remarks. In addition to the type species, material representing three un-
named species was studied in an evaluation of the generic characters.
Kethley: Syringophilidae 25
Ascetomylla gallinula, new species
(Figs. 12, 12)
FEMALE (holotype). Length 1380u; width 345u. Gnathosoma: Hypostomal
apices (Fig. 12 C, D), highly ornate, seven pairs of processes, of which the
outer three pairs are longest. Peritremes with each lateral branch three cham-
bered; each longitudinal branch with 18-19chambers. Dorsal idiosoma: Pro-
podosomal plate divided, bearing setae vi, ve; ratios of vi:ve:sci:sce, 1:3:3:4;
sci, dl, 11 subequal. Hysterosomal plate absent; 12, d3, 13 subequal to sce;
ratios of d4:14:d5:15, 8:8:1:8; d4 1 1/2 times length of dl. Ventral idiosoma:
As in Fig. 12 B. Legs: Setae dF, dG, QT of legs I and II smooth; a’ and a" I-IV
each with 15-16 tines, a'I 1/2 length of a"I; sc3 and sc4 subequal, not extend-
ing beyond genua; vFII extending at least to ambulacrum.
MALE (paratype). Length 980u; width 320u. Gnathosoma: Hypostomal
apices unornamented. Peritremes as in female. Dorsal idiosoma: Propodosomal
plate not divided, bearing setae vi, ve, dl; ratios of vi:ve:sci:sce, 1:1:1:2; 11
2/3 length of sce; dl subequal to sce. Hysterosomal plate absent; 12, d3, 13
subequal, 1/2 length of vi; ratios of d4: 14:gl:g2, 2:6:1:1; d4 subequal to vi.
Ventral idiosoma and aedeagus: As in Fig. 13 B. Legs: As in female except
a'I subequal to a"l.
Type material. From Gallinula chloropus (L.), Rallidae: holotype female;
2 male and 2 female paratypes Johannesburg, Transvaal, Union of South Africa;
February 9, 1954; J. Gaud. The holotype and one paratype male and one para-
type female are deposited with the South African Institute for Medical Research.
The remaining paratypes are deposited with the Entomology Department, Univer-
sity of Georgia, Athens, Georgia. The name gallinula refers to the host.
Trypetoptila, new genus
This new genus appears most similar to Syringophilopsis. Species of Try-
petoptila may be distinguished by the presence of 3-6 additional pairs of setae
in the paragenital series, and the presence of leg setae vs'III. These very large
mites (1520u- 1880) are found in the covert feathers of ciconiiform birds of
the family Ardeidae. The name Trypetoptila (feminine singular) means feather
borer and refers to the hole found in the quills inhabited by the mite.
Type species: Trypetoptila casmerodia, new species, ex. Casmerodius
albus egretta (Ord); Ardeidae, Ciconiiformes.
FEMALE. (1). Hypostomal apex ornamented; rounded protuberances present.
(2). Lateral hypostomal teeth absent. (3). Chelicerae dentate, 2-3 teeth. (4).
Peritreme M shaped; lateral branches each with 2-3 chambers, longitudinal
branches each with 8-10 chambers. (5). Stylophore rounded posteriorly, not
extending below propodosomal plate. (6). Palpal tibiotarsus rounded on distal
margin. (7). All setae smooth. (8). Propodosomal plate cleft; lateral margin
concave. (9). Hysterosomal plate absent. (10). Setal pattern of propodosomal
region with six pairs of setae arranged 3-1-2. (11). Setae 12, d3, 13 long; d3
closer to12 than to13. (12). Setae d4, 14, d5, 15 long. (13). Genital series
with two pairs of setae; anal series with two pairs of setae. (14). Paragenital
series supernumerary; 6-9 pairs of setae present. (15). MCAI1 divergent, fused
to MCA2 at anterior 1/3 to 1/2 length of MCA2. (16). Coxae III and IV weakly
sclerotized, margins indistinct. (17). Cuticular striations as in Fig. 14A, B.
(18). Legs I-IV subequal in thickness. (19). Legs with supernumerary setal
26
Contrib. Amer. Ent. Inst., vol. 5, no. 6, 1970
lOOu
s
es
en
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f
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<a
7
LL]
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cA
oN
A
ae
4 3 \
Female (holotype): A,
dorsal aspect; B, ventral aspect; C, hypostomal lips, dorsal aspect;
D, hypostomal lips, ventral aspect.
Fig. 12. Ascetomylla gallinula new species.
Kethley: Syringophilidae 27
Fig. 13. Ascetomylla gallinula new species. Male (paratype): A,
dorsal aspect; B, ventral aspect.
condition, vs'III present. (20). Setae a' anda" multiserrate, 24-28 tines.
(21). Antaxial and paraxial members of claw pair subequal with basal angles;
claws of equal length as empodium. (22). Order of host: Ciconiiformes.
(23). Types of feathers inhabit: secondaries and coverts.
MALE. As in female except: (1). Hypostomal lips absent. (3). Chelicerae
edentate. (8). Propodosomal plate weakly sclerotized, margins indistinct. (10).
Setal pattern of propodosomal region 3-2-1. (11). Setae d3, 13 short. (12). Setae
d4,d05,15 short. (14). Paragenital series supernumerary, with 5-6 pairs of setae
present. (17). Cuticular striations as in Fig. 15A,B. (21). Claws 1/3 length of
empodium.
28 Contrib. Amer, Ent. Inst. , vol:5; no. 6, 1970
Trypetoptila casmerodia, new species
(Figs. 14, 15)
FEMALE (holotype). Length 1880u; width 320u. Gnathosoma: Hypostomal
apices slightly ornamented, two pairs of blunt processes. Peritremes (Fig.14 C),
with each lateral branch 3 chambered; each longitudinal branch 8 chambered.
Dorsal idiosoma: Propodosomal plate not divided, concave on posterior margin,
bearing setae vi, ve, sci, dl, 1]; ratios of vi:ve:sci:sce, 1:1:11/4:11/4; sce,
dl, 11 subequal. Hysterosomal plate absent; 12, d3, 13 slightly longer than dl;
ratio of d4:14:d5:15, 1:1:1:1; d4 subequal to dl. Ventral idiosoma: As in Fig. 14
B. Legs: Setae dF, dG, dT of legs I, II smooth; a' and a" I-IV with 22-26 tines
each, a'I 1/2 length of a"I; sc3 and sc4 subequal, not extending beyond genua;
vFII extending at least to ambulacrum.
MALE (paratype). Length 1260uU; width 410u. Gnathosoma: Hypostomal
apices unornamented. Peritremes as in female. Dorsal idiosoma: Propodosomal
plate not divided, bearing setae vi, ve, sci, dl, 11; ratios of vi:ve:sci:sce,
1:1:11/2:2; sci subequal to dl, 11. Hysterosomal plate absent; 12 subequal to
sce; d3, 13 1/2 length 12; ratio of d4:14:gl:g2, 3:6:1:1; d4 subequal to d3. Ventral
idiosoma and aedeagus: As in Fig. 15 B. Legs: As in female except a'I sub-
equal to a'Il.
Type material. From Casmerodius albus egretta (Ord), Ardeidae; holotype
female, 7 male and 18 female paratypes, McIntosh County, Georgia, March 19,
1969; W. T. Atyeo, C. W. Proctor, and J. B. Kethley. The holotype and para-
types are deposited with the Entomology Department, University of Georgia,
Athens, Georgia. The name casmerodia refers to the host.
Torotrogla, new genus
Among those genera having MCA] strongly divergent and fused to MCA2
(Torotrogla, Selenonycha and Creagonycha), Torotrogla is distinguished by the
presence of 1-3 additional pairs of setae in the paragenital series. The affin-
ities of this new genus are uncertain. These medium sized to large mites (640u-
890) are found in the primaries of passeriform birds of the families Fringillidae,
Mimidae, Ptilgonatidae, and Turdidae. The name Torotrogla (feminine, singular)
means hole borer and refers to the hole found in the feathers inhabited bythe mites.
Type species: Torotrogla mima, new species, ex. Mimus polyglottos (Lo
Mimidae, Passeriformes. :
FEMALE. (1). Hypostomal apex ornamented; paired median protuberance
present. (2). Lateral hypostomal teeth absent. (3). Chelicerae dentate, 2-3
teeth. (4). Peritreme M shaped; each lateral branch with 2-3 chambers; each
longitudinal branch with 4-12 chambers. (5). Stylophore rounded posteriorly,
not extending beneath propodosomal plate. (6). Palpal tibiotarsus rounded on
distal margin. (7). All setae smooth. (8). Propodosomal plate variable, weakly
sclerotized; usually rectangular. (9). Hysterosomal plate variable. (10). Setal
pattern of propodosomal region with six pairs of setae arranged 3-2-1. (11). Setae
12, d3, 13 long; d3 closer to 13 than to12. (12). Setae d4, 14 long, d5, 15 short.
(13). Genital series with two pairs of setae; anal series with two pairs of setae.
(14), Paragenital series supernumerary; 4-6 pairs of setae present. (15). MCAI
strongly divergent, fused to MCA2 at basal 1/2 length of MCA2. (16). Coxae III
and IV weakly sclerotized, margins indistinct. (17). Cuticular striations as in
Fig. 16 A, B. (18). Legs I slightly thicker than II; legs II - IV subequal in
Kethley: Syringophilidae 29
Fig. 14. Trypetoptila casmerodia new species. Female (holotype): A,
dorsal aspect; B, ventral aspect; C. peritreme.
thickness. (19). Legs with full complement of setae. (20). Setae a' and a"
multiserrate, 10-20 tines. (21). Antaxial and paraxial members of claw pair
subequal; claws extending 1/5 length of empodium. (22). Order of hosts:
Passeriformes. (23). Types of feathers inhabit: primaries.
MALE. As in the female except: (1). Hypostomal lips absent. (2). Cheli-
cerae edentate. (5). Stylophore rounded posteriorly or constricted. (11). Setae
12, d3, 13 short. (12). Setae d4 short. (14). Paragenital series with 3-4 pairs
of setae. (17). Cuticular striations as in Fig. 17A, B. (21). Claws 1/3 length
of empodium.
30 Contrib. Amer. Ent. Inst., vol. 5, no. 6, 1970
100p Ai —=B
aa
- TANS
eioree ls ee
1p!
@) if © e) GQ @e
‘y ae QO!
\ Za
© G \
a \P-- IRE
®) @ WE !
G) A !
D No uf Q
N.
G) SN.
> sav’ G)
Fig. 15. Trypetoptila casmerodia new species. Male (paratype): A,
dorsal aspect; B, ventral aspect.
The genus Torotrogla includes one named and 5 new species:
Torotrogla villosa (Hancock)
new combination
Picobia villosa Hancock, 1895, Amer. Nat. 29:382-384, Fig. 1,2.
All type material of this species is lost. Originally described from Phaino-
pepla nitens (Swainson) (Passeriformes: Ptilgonatidae) in Arizona, this species
has not been recollected.
Kethley: Syringophilidae 31
Torotrogla mima, new species
(Figs. 16. 17)
FEMALE (holotype). Length 890u; width 300. Gnathosoma: Hypostomal
apices ornamented, with paired median protuberances. Peritremes (Fig. 16 C),
each lateral branch 3 chambered; each longitudinal branch 6 chambered. Dorsal
idiosoma: Propodosomal plate not divided, bearing setae vi, ve, sci, dl; ratios
of vi:ve:sci:sce, 1:11/2:3:4; sce, dl, 11 subequal in length. Hysterosomal
plate absent; 12, 13 subequal in length to dl; d3 three times length of vi; ratios
of d4:14:d5:15, 7:7:1:1; d4 two times length of dl. Ventral idiosoma: As in Fig.
16 B. Legs: Setae dF, dG, dT of legs I and II smooth; a‘ and a" I-IV each with
16-17 tines, a'l 2/3 length of a"I; sc3 and sc4 subequal, extending beyond
genua; vFII extending at least to ambulacrum; tc'III and IV each subequal to
te "III and IV.
MALE (paratype). Length 740u; width 280uU. Gnathosoma: Hypostomal
apices unornamented. Peritremes as in female. Dorsal idiosoma: Propodosomal
plate not divided, bearing setae vi, ve, sci, dl; ratios of vi:ve:sci:sce, 1:1:3:4;
dl, 11 four times length of vis Hysterosomal plate present and continuous with
pygidial plate, bearing setae d3, d4, 14, d5, 15; 12, d3, 13, vi subequal in
length; ratios of d4:1l4:glig2, 2:10:1:1; d4 subequal to sci; pg3 dorsal. Ventral
idiosoma and aedeagus: As in Fig. 17 B. Legs: As in female except a'I subequal
to a"I; sc3 and sc4 not extending beyond genua.
Type material. From Mimus polyglottos (L.), Mimidae: holotype female,
thirty female and fourteen male paratypes, one mile N. W. Odum, Wayne County,
Georgia, March 20, 1969, W.T. Atyeo, C.W. Proctor, J. B. Kethley. Holotype,
fifteen female and seven male paratypes deposited with the University of Georgia,
Athens, Georgia.
The name mima refers to the host.
Selenonycha, new genus
Within the group of genera having MCAl1 strongly divergent, Selenonycha is
distinguished by the broadly open shape of the claws of the adult female and the
presence of three pairs of paragenital setae. This new genus appears most sim-
ilar to Creagonycha. These large mites (930U- 1360) are found in the flight
feathers of charadriform birds of the families Charadriidae, Laridae, and Scolo-
pacidae. The name Selenonycha (feminine singular) means crescent shaped claws
and refers to the shape of the claws of the adult female.
Type species: Selenonycha baltoda, new species, ex. Charadrius wilsonia
wilsonia Ord,; Charadriidae, Charadriiformes.
FEMALE. (1). Hypostomal apex ornamented slightly. (2). Lateral hypo-
stomal teeth absent. (3). Chelicerae with three teeth. (4). Peritreme M-shaped;
lateral branches each with 4-6 chambers; longitudinal branches each with 8-12
chambers. (5). Stylophore rounded posteriorly; extending to anterior margin of
propodosomal plate. (6). Palpal tibiotarsus rounded on distal margin. (7). All
setae smooth. (8). Propodosomal plate weakly sclerotized, cleft; lateral margins
of plate concave; anterior margin concave; 11 not on plate. (9). Hysterosomal
plate variable. (10). Setal pattern of propodosomal region with six pairs of
setae arranged 3-2-1 to 2-3-1 due to variable nature of 11. (11). Seate 12, 13,
d3 long; d3 closer to 12 than to13. (12). Seate d4, 14, d5, 15 long; d4 lateral;
d5 on pygidial plate. (13). Genital series with two pairs of setae; anal series
32 Contrib. Amér. Ent. Inst. , vol. 5; no. 6, 1970
lOOu
Fig. 16. Torotrogla mima new species. Female (holotype): A, dorsal
aspect; B, ventral aspect; C, peritreme.
with two pairs of setae. (14). Paragenital series with three pairs of setae. (15).
MCA] strongly divergent, fused to MCA2; MCA] similar in shape to MCA2. (16).
Coxae III and IV weakly sclerotized, margins indistinct. (17). Cuticular stria-
tions as in Fig. 18 A, B. (18). Legs I-IV subequal in thickness. (19). Legs with
full complement of setae. (20). Setae a' and a" multiserrate, 18-26 tines. (21).
Antaxial and paraxial members of claw pair subequal; claws broadly open in shape;
Kethley: Syringophilidae 38
Pig. 17. Torotrogla mima new species. Male (paratype): A, dorsal
aspect; B, ventral aspect.
extending to full length of empodium. (22). Order of hosts:
(23). Types of feathers inhabit: unknown.
MALE. As in female except: (1). Hypostomal hyaline lips lacking. (3). Cheli-
cerae edentate. (11). Setae13, d3 short. (12). Setae d4, d5, 15 short. (17).
Cuticular striation patterns as in Fig. 19A,B. (21). Claws 1/2 length of empodium.
In addition to the type species, material representing two unnamed species
was studied in an evaluation of the generic characters.
Charadriiformes.
34 Contrib. Amer: Ent,:.Inst.; vol. 5,.no. 6,°1970
Selenonycha baltoda, new species
(Figs. 18, 19)
FEMALE (holotype). Length 1070u; width 312u. Gnathosoma: Hypostomal
apices unornamented, rounded. Peritremes with each lateral branch 5 chambered:
each longitudinal branch 8 chambered. Dorsal idiosoma: Propodosomal plate
not divided, cleft on posterior margin, lateral margins concave, bearing setae
vi, ve, dl; ratios of vi:ve:sci:sce, 1:4:8:8; dl, 11, sce subequal. Hysterosomal
plate absent; 12, d3, 13 subequal, each 11/2 length of ve; ratios of d4:14:d5:15,
l:1:1:1; 14, three times length of ve. Ventral idiosoma: As in Fig. 18 B. Legs:
Setae dF, dG, aT of legs I and II smooth; a' and a" I-IV each with 14-15 tines;
a'I subequal to a"I; sc3 and sc4 subequal, extending beyond genua; vFI, II sub-
equal, each not extending to ambulacrum; tc'III and IV subequal 2/3 length of
te Tr and Iv;
MALE (paratype). Length 840u; width 300u. Gnathosoma: Hypostomal
apices unornamented. Peritremes as in female. Dorsal idiosoma: Propodosomal
plate not divided, bearing setae vi, ve, sci, 11; ratios of vi:ve:sci:sce, 1:2:3:4;
11 subequal to sce; dlsubequal to vi. Hysterosomal plate absent; 12subequal to
sci; d3, 13 each subequal to vi; ratios of d4:14:gl:g2, 2:16:1:1; 14 twice length
of sce. Ventral idiosoma and aedeagus: As in Fig. 19 B. Legs: As in female
except a'l1/2 length of a"I; sc3, sc4 not extending beyond genua; tc'III and IV
1/2 length of tc"III and IV.
Type material. From Charadrius wilsonia wilsonia Ord, Charadriidae;
holotype female, 4 female paratypes, 1 male paratype, Cameron Parish, Louisiana,
June, 1966, R. Heard. The holotype and paratypes are deposited with the Uni-
versity of Georgia, Athens, Georgia.
The name baltoda refers to the habitat of the host.
Creagonycha, new genus
The strongly recurved shape of the claws of species of Creagonycha dis-
tinguishes this genus from other genera having MCAI strongly divergent and
fused to MCA2. This new genus appears most similar to Selenonycha. These
medium sized to large mites (740U- 975) are found in the secondaries of cha-
radriiform birds of the families Scolopacidae and Laridae. The name Creagonycha
(feminine singular) means hooked claw and refers to the condition of the claw of
the adult female.
Type species: Creagonycha lara, new species; ex. Larus delawarensis Ord,
1815; Laridae, Charadriiformes.
FEMALE. (1). Hypostomal apex slightly ornamented. (2). Lateral hypostomal
teeth absent. (3). Chelicerae with three teeth. (4). Peritreme M-shaped; lateral
branches each 4-5 chambered; longitudinal branches each with 14-18 chambers.
(5). Stylophore rounded on posterior margin, not extending below propodosomal
plate. (6). Palpal tibiotarsus rounded on distal margin. (7). All setae smooth.
(8). Propodosomal plate weak sclerotized, all margins concave; 11, dl not on
plate. (9). Hysterosomal plate variable. (10). Setal pattern of propodosomal
region with six pairs of setae arranged 3-2-1. (11). Setae 12, d3, 13 long; d3
closer to 12 than to13. (12). Setae 14, d5, 15 long, subequal in length; d4 short,
length variable from 1/8 to 1/2 of length of 14. (13). Genital series with two
pairs of setae; anal series with two pairs of setae. (14). Paragenital series with
three pairs of setae. (15). MCAlI strongly divergent, fused to MCA2; MCAI1 and
Kethley: Syringophilidae 35
ihe,
a Ns
ie
Pe cnNes aoe ae
Fig. 18. Selenonycha baltoda new species. Female (holotype): A,
dorsal aspect; B, ventral aspect; C,claw, tarsus IV.
MCA2 similar in shape. (16). Coxae III and IV weakly sclerotized, margins in-
distinct. (17). Cuticular striations as in Fig. 20 A, B. (18). Legs I-IV subequal
in thickness. (19). Legs with full complement of setae. (20). Setae a' anda"
multiserrate, 14-22 tines. (21). Antaxial and paraxial members of claw pair
subequal, strongly recurved in shape; (Fig. 20 C) claws 1/2 length of empodium.
(22). Order of hosts: Charadriiformes. (23). Types of feathers inhabit: secon-
daries and coverts.
MALE. As in female except: (1). Hypostomal lips without hyaline lips. (3).
Chelicerae edentate. (8). Propodosomal plate weakly sclerotized, all margins
concave; 11 not on plate. (11). Setae 13, d3 short. (12). Setae 14, d4, 15 short.
(17). Claws recurved, but not as pronounced as in female; extending 1/3 length of
empodium.
The genus Creagonycha includes one named and 4 new species.
36 Contrib. Amer. ent. Inst:, VOl-5> no. 6, 1970
si Si,
PIN Ss
ee
ae SN
; ports \
( x XN a
SoeNG N
Roe NX
N
=
GGL
Va
Fig. 19. Selenonycha baltoda new species. Male (paratype): A,
dorsal aspect; B, ventral aspect.
Creagonycha lara, new species
(Figs. 20, 21)
FEMALE (holotype). Length 1060u; width 330u. Gnathosoma: Hypostomal
apices unornamented. Peritremes, each lateral branch one chambered; each lon-
gitudinal branch 15 chambered. Dorsal idiosoma: Propodosomal plate not di-
vided , bearing setae vi, ve, sci; ratios of vi:ve:sci:sce, 1:2:6:61/2; sci, dl, 11
subequal in length to dl; ratios of d4:14:d5:15, 3:4:3:2; d4 subequal to length of
dl. Ventral idiosoma: As in Fig 20 B. Legs: Setae dF, dG, dT of legs I and Il
smooth; a' and a" with 14-15 tines each, a'I subequal to a"I; sc3 and sc4 sub-
equal, not extending beyond genua; vFII extending at least to ambulacrum.
Kethley: Syringophilidae 37
MALE (paratype). Length 860u; width 305u. Gnathosoma: Hypostomal apices
unornamented. Peritremes as in female. Dorsal idiosoma: Propodosomal plate
not divided, bearing setae vi, ve, sci, dl; ratios of vi:ve:sci:sce, 1:1:2:3; dl,
11 each 31/4 times length of vi. Hysterosomal plate absent; 12 2/3 dl; d3 1/2
dl; 13 1/3 dl; ratios of d4:14:gl:g2, 3:9:1:1; d4 subequal to vi. Ventral idiosoma
and aedeagus: As in Fig. 21 B. Legs: As in female except a'I subequal to a'II.
Type material. From Larus delawarensis Ord, Laridae: holotype female, 6
female paratypes, 3 male paratypes, Coffee Bluff, Chatham County, Georgia, March
18, 1969, W.T. Atyeo, C.W. Proctor, J. B. Kethley. Holotype and all paratype
deposited with the University of Georgia, Athens, Georgia.
The name lara refers to the host.
Creagonycha totana (Oudemans)
new combination
Syringophilus totani Oudemans, 1904, Entomol. Ber., 19:171; 1906, Mem. Soc.
4001, Paris-19) S/-62,.Fige. 7, @,
The holotype, on loan from Laboratoire d'Acarology, was examined. De-
scribed originally from Tringa totanus (L.) (=Totanus calidris = Totanus totanus)
(Charadriiformes: Scolopacidae), in France, this species has not been recollected.
Chenophila, new genus
Among those genera having a posterior constriction of the stylophore (Chen-
ophila, Stibarokra and Syringonomus), Chenophila is distinguished by the arrange-
ment of the setae of the propodosomal region and the presence of the full com-
plement of leg setae. The affinities of this new genus are difficult to ascertain,
as the condition of the stylophore indicates only similarity rather than relation-
ship. These medium sized mites (750uU- 820) are found in the flight feathers of
anseriform birds of the family Anatidae. The name Chenophila (feminine sing-
ular) means goose lover and refers to the host relationship.
Type species: Chenophila branta, new species; ex. Branta canadensis (L.);
Anatidae, Anseriformes.
FEMALE. (1). Hypostomal apex unornamented. (2). Lateral hypostomal teeth
absent. (3). Chelicerae with one tooth. (4). Peritreme M-shaped; lateral branches
each with 14-16 chambers. (5). Stylophore constricted posteriorly; subcutaneous
to propodosomal plate to level of sci. (6). Palpal tibiotarsus rounded on distal
margin. (7). All setae smooth. (8). Propodosomal plate entire, strongly scle-
rotized, rectangular. (9). Hysterosomal shield variable. (10). Setal pattern of
propodosomal region with six pairs of setae arranged 3-1-2. (11). Setae 12, d3,
13 long; d3 closer to_12 than13, or equidistant between 12 and13. (12). Seta 14
long; d4, d5, 15 short; 14 on terminal tubercle. (13). Genital series with two
pairs of setae; anal series with two pairs of setae. (14). Paragenital series
with three pairs of setae. (15). MCAI1 weakly divergent; fusion of MCAl to MCA2
indistinct. (16). Coxae III and IV strongly sclerotized, rectangular in shape.
(17). Cuticular striations as in Fig. 28 A, B; longitudinal between coxae III and
IV. (18). Legs I thicker than II and legs II thicker than III and IV. (19). Legs
with full complement of setae. (20). Setae a' anda" multiserrate, 6-10 tines.
(21). Antaxial and paraxial member of claw pair subequal; claws 1/2 length of
empodium. (22). Order of hosts; Anseriformes. (23). Types of feathers in-
habit: coverts. Only the female is known.
In addition to the type species, material representing twelve unnamed species
was examined in an evaluation of the generic characters.
38 Contrib. Amer. Ent. Inst., vol. 5, no. 6, 1970
Fig. 20. Creagonycha lara new species. Female (holotype): A,
dorsal aspect; B, ventral aspect; C, claw, tarsus IV.
Chenophila branta, new species
(Fig. 22)
FEMALE (holotype). Length 810u; width 218u. Gnathosoma: Hypostomal
apices unornamented, smooth. Peritremes (Fig. 22 E) with each lateral branch
five chambered; each longitudinal branch 14 chambered. Dorsal idiosoma: Propo-
dosomal plate not divided, bearing setae vi, ve, sci, dl, 11; ratios of vi:ve:sci:
Kethely: Syringophilidae 39
Pig. 21. Creagonycha lara new species. Male (paratype): A, dorsal
aspect; B, ventral aspect.
sce, 1:11/2:13/4:2 1/4; dl, 11, sce subequal. Hysterosomal plate absent; 12,
d3, 13 subequal, each 1/2 length of sce. Ventral idiosoma: As in Fig. 22 B.
gs: Setae dF, dG, dT of legs I and II smooth; a' and a" I-IV each with 12-13
tines; a'I 1/2 length of a"I; sc3 and sc4 subequal, not extending beyond genua;
vF I and II subequal, extending at least to ambulacrum; tc'III and IV subequal,
2/3 length of tc"III and IV. Only the female is known.
Type material. From Branta canadensis (L.), Anatidae: holotype female,
40 Contrib. Amer. Ent. Inst,,:vol..5, no. 6,.1970
\OOu
Fig. 22. Chenophila branta new species. Female (holotype): A, dorsal
aspect; B, ventral aspect; C, hypostomal lips, dorsal aspect; D, hypo-
stomal lips, ventral aspect.
Kethley: Syringophilidae 4l
30 female paratype, Kensington Park, Oakland County, Michigan, July 6, 1967,
J.B. Kethley. The holotype and 10 female paratypes are deposited at the Uni-
versity of Georgia, Athens, Georgia.
The name branta refers to the host.
Aulobia, new genus
Among those genera having parallel MCA1, (Aulobia, Aulonastus, Niglarobia,
Syringophiloidus, and Philoxanthornea), Aulobia is distinguished by the presence
of the full complement of leg setae, and the lack of a basal angle of the claws.
This genus appears to stand as a transition group between those genera of large
mites with curved and fused MCA] and those genera of small mites with parallel
MCA] and reduced leg setation. These small to medium sized mites (650U- 800)
are found in the flight feathers of passeriform birds of the families Parulidae and
Sylviidae. The name Aulobia (masculine singular) means inhabitant of small
quills, and refers to the avian hosts.
Type species: Syringophilus dendroicae Clark, 1964, Acarologia 6: 86, Fig.
21; ex. Dendroica coronata (L.); Parulidae, Passeriformes.
FEMALE. (1). Hypostomal apex slightly ornamented, one pair median pro-
tuberances present. (2). Lateral hypostomal teeth absent. (3). Chelicerae eden-
tate. (4). Peritreme M-shaped lateral branches each with 2-4 chambers; longi-
branches each with 10-16 chambers. (5). Stylophore rounded, not extending be-
low propodosomal plate. (6). Palpal tibiotarsus rounded on distal margin. (7).
All setae smooth. (8). Propodosomal plate entire; lateral margins parallel; ante-
rior margin concave; posterior margin scalloped. (9). Hysterosomal plate vari-
able. (10). Setal pattern of propodosomal region with six pairs of setae arranged
2-1-12. (11). Setae 12, 13, d3 long; d3 closer to_12 than to13. (12). Seate d4,
14 long; d5, 15 short. (13). Genital series with two pairs of setae; anal series
with two pairs of setae. (14). Paragenital series with three pairs of setae. (15).
MCAI parallel, not fused to MCA2. (16). Coxae III and IV rectangular, subequal,
strongly sclerotized. (17). Cuticular striation patterns as in Fig. 23 A, B. (18).
Legs I-IV subequal in thickness. (19). Legs with full complement of setae. (20).
Setae a' and a" multiserrate; 6-10 tines. (21). Antaxial and paraxial members of
claw pair subequal; claws 1/3 length of empodium. (22). Order of hosts: Passeri-
formes. (23) Types of feathers inhabit: primaries and secondaries.
MALE. As in female except: (1). Hypostomal apex unornamented. (11). Setae
12,13, d3 short. (15). MCAl waekly divergent, free, not fused to MCA2. (16).
Coxae III and IV weakly sclerotized, margins indistinct. (17). Cuticular stria-
tions as in Fig. 24 A, B.
The genus Aulobia includes one named and two new species. The named
species transferred to this genus is:
Aulobia dendroicus (Clark)
new combination
(Pies, 23,24)
Syringophilus dendroicae Clark, 1964, Acarologia 6: 86, Fig. 21.
Although the holotype is lost, thrity-two paratypes on loan from the Public
Health Service Rocky Mountain Laboratory were examined. Described originally
from Dendroica coronata (L.) (Passeriformes: Parulidae), in Maryland, this species
has been collected from the same host in Florida and Georgia, U. S.A.
42 Contrib. Amer. Ent. Inst., vol. 5, no. 6, 1970
|OOu
Fig. 23. Aulobia dendroicus (Clark). Female (paratype): A, dorsal aspect;
B. ventral aspect; C, hypostomal lips, dorsal aspect; D, hypostomal lips,
ventral aspect.
Kethley: Syringophilidae 43
Fig. 24. Aulobia dendroicus (Clark). Male (paratype): A, dorsal aspect;
B, ventral aspect.
Niglarobia, new genus
Within the previously mentioned genera having MCA] parallel, Niglarobia is
distinguished by the arrangement of the setae of the propodosomal region, claws
with a basal angle, the loss of leg setae vs'II. This new genus is similar to
Aulobia, but differs in the reduced leg setation. These small to medium sized
mites (520U- 650) are found in the primaries and secondaries of charadriiform
birds of the family Scolopacidae. The name Niglaobia (masculine plural) means
44 Contrib, Amer. Ent. Insts; vols 5.-no. 6; 1970
inhabitant of small quills and refers to the host.
Type species: Niglarobia ereuneti, new species; ex. Ereunetes pusillus
(L.); Scolopacidae, Charadriiformes.
FEMALE. (1). Hypostomal apex unornamented. (2). Lateral hypostomal teeth
absent. (3). Chelicerae edentate. (4). Peritreme M-shaped; lateral branches
each with 2-4 chambers; longitudinal branches each with 6-10 chambers. (5).
Stylophore rounded posteriorly, extending to edge of propodosomal plate. (6).
Palpal tibiotarsus rounded on distal margin. (7). All setae smooth. (8). Propo-
dosomal plate weakly sclerotized, margins indistinct; lateral margins parallel;
anterior margin concave; posterior margin scalloped. (9). Hysterosomal plate
variable. (10) . Setal pattern of propodosomal region with six pairs of setae pat-
tern of propodosomal region with six pairs of setae arranged 2-1-1-2. (11). Setae
12, 13, d3 long; d3 closer to_12 than to13. (12). Setae d4, 14 long, d4 1/2 length
of 14; dS, 15 short; d5 on pygidial plate. (13). Genital series with two pairs of
setae; anal series with two pairs of setae. (14). Paragenital series with three
pairs of setae. (15). MCAlI parallel, not fused to MCA2. (16). Coxae III and IV
rectangular in shape, weakly sclerotized, margins indistinct; articulation to
trochanters prominent. (17). Cuticular striations as in Fig. 25 A,B. (18). Legs
I-IV subequal in thickness. (19). Setae vs'II absent. (20). Setae a' anda".
multiserrate, 6-10 tines. (21). Antaxial and paraxial members of claw pair sub-
equal, with basal angle; claws 1/3 length of empodium. (22). Order of hosts:
Charadriiformes. (23). Types of feathers inhabit: secondaries and primaries.
MALE. As in female except: (8). Propodosomal plate weakly sclerotized,
margins indistinct; dl, 11 not on plate. (10). Setal pattern of propodosomal
region with six pairs of setae arranged 2-1-2-1. (11). Setae 13, d3 short. (12).
Setae d4 short. (17). Cuticular striations as in Fig. 26 A, B.
The genus Niglarobia includes two named and four new species.
Niglarobia ereuneti, new species
(Pigs. 25, 26)
FEMALE (holotype). Length 490u; width 115u. Gnathosoma: Hypostomal
apices unornamented, smooth. Peritremes with each lateral branch one cham-
bered; each longitudinal branch 6 chambered. Dorsal idiosoma: Propodosomal
plate not divided, bearing setae vi, ve, sci, dl, 11; ratios of vi:ve:sci:sce,
1:1:11/4:4; sce, dl, 11 subequal in length. Hysterosomal plate absent; 12, d3
three times length of vi; 13 five times length of dl. Ventral idiosoma: As in Fig.
25 B. Legs: Setae dF, dG, dT of legs I and II smooth; a' and a" I-IV with six
tines each, a'I 1/2 length of a"I; sc3 and sc4 subequal, extending beyond genua;
vFII extending at least to ambulacrum; tc'III and IV subequal each 1/2 length of
te"IIT and IV.
MALE (paratype). Length 400u; width 120u. Gnathosoma: Hypostomal
apices unornamented. Peritremes as in female. Dorsal idiosoma: Propodosomal
plate not divided, bearing setae vi, ve, sci, dl; ratios of vi:ve:sci:sce, 1:1:2:6;
dl, 11, sce subequal. Hysterosomal plate absent; d3, 13 subequal to vi; 12 twice
length of vi; ratios of d4;l4:gl:gq2, 2:8:1:1; d4 twice length of vi. Ventral idio-
soma and aedeagus: As in Fig. 26 B. Legs: As in female except a'I subequal to
a'Il; sc3 and sc4 subequal, not extending beyond genua.
Type material. From Ereunetes pusillus (L.), Scolopacidae: holotype female,
1 male and 8 female paratypes, Sapelo Island, Georgia, February 12,1968, W. B.
Sikora. Holotype, 1 male and 4 female paratypes deposited with University of
Kethley: Syringophilidae 45
lOOu
Fig. 25. Niglarobia ereuneti new species. Female (holotype): A, dorsal
aspect; B, ventral aspect; C, claw, tarsus IV.
46 Contrib. Amer. Ent. Inst., vol.
Fig. 26. Niglarobia ereuneti new species.
aspect; B, ventral aspect.
Georgia, Athens, Georgia.
The name ereuneti refers to the host.
fi ne.-0, 2970 —
Male (paratype): A, dorsal
Kethley: Syringophilidae 47
Niglarobia helleri (Oudemans)
new combination
Syringophilus helleri Oudemans, 1904, Entomol. Ber. 20:190; 1906, Mem. Soc.
zool. Paris 19:62-66, Figs. 9, 1l.
The holotype of this species on loan from the Laboratoire d'Acarologie was
examined. Described originally from Tringa ochropus (L.) (=Totanus ochropus)
(Charadriiformes: Scolopacidae), in France, this species has not beenrecollected.
Niglarobia trouessarti (Oudemans)
new combination
Syringophilus troussarti Oudemans, 1904, Entomol. Ber. 20:190; 1906, Mem.
Soc. zool. Paris 19:67-72, Figs. 12-15.
The holotype of this species on loan from the Laboratoire d'Acarologie was
examined. Described originally from Aramus scolopaceus (Gmelin) (Charadriifor-
mes: Scolopacidae), in South America, this species has not been recollected.
Aulonastus, new genus
Among those genera having MCAI parallel and not fused to MCA2, Aulonastus
is distinguished by the loss of the following setae: vi, dTIII, dIIV, and a pair
of anal setae. These small mites (425u- 500y) are found in the flight feathers of
passeriform birds of the family Fringillidae and piciform birds of the family Picidae.
The name Aulonastus (masculine plural) means dwelling in a small tube and refers
to the avian hosts.
Type species: Aulonastus pipili, new species; ex. Pipilo erythrophthalmus
(L.); Fringillidae, Passeriformes.
FEMALE. (1). Hypostomal apex slightly ornamented; paired median protu-
berances present. (2). Lateral hypostomal teeth absent. (3). Chelicerae edentate.
(4). Peritreme M-shaped; each lateral branch 2-3 chambered; each longitudinal
branch with 6-8 chambers. (5). Stylophore rounded posteriorly, not extending
beneath propodosomal plate. (6). Palpal tibiotarsus rounded on distal margin.
(7). All setae smooth. (8). Propodosomal plate weakly sclerotized, margins in-
distinct; rectangular in shape. (9). Hysterosomal plate variable. (10). Setal
pattern of propodosomal region with five pairs (vi absent) arranged 1-1-1-2. (11).
Setae 12 variable, long or short; 13, d3, short; d3 closer to12 thanto13. (12).
Setae 14 long; d4, d5, 15 short; 14 on tubercle; 15 on pygidial plate. (13). Genital
series with two pairs of setae; anal series with one pair of setae. (14). Para-
genital series with three pairs of setae. (15). MCAI parallel, not fused to MCA2.
(16). Coxae III and IV weakly sclerotized, margins indistinct; apophysis of artic-
ulation to trochanters prominent. (17). Cuticular striations as in Fig. 27 A, B.
(18). Legs I thicker than II, and II thicker than III and IV. (19). Leg setae scl,
sc2, QTIII, dTIV absent. (20). Setae a' and a" multiserrate, 6-8 tines. (21).
Antaxial and paraxial members of claw pair subequal; claws 1/3 length of empo-
dium. (22). Orders of hosts: Passeriformes and Piciformes. (23). Types of feath-
ers inhabit: coverts.
MALE. Unknown.
48 Contrib. Amer. Ent. Inst., vol. 5, no. 6, 1970
Aulonastus pipili, new species
(Fig. 27)
FEMALE (holotype). Length 470uU; width 120uU. Gnathosoma: Hypostomal
apices unornamented, smooth. Peritremes with each lateral branch one cham-
bered; each longitudinal branch 6 chambered. Dorsal idiosoma: Propodosomal
plate not divided, bearing setae ve, sci, dl, 11; vi absent; ratios of ve:sci:sce:
di, T:istiitis el and 11 subequal. Hysterosomal plate absent; 12 subequal to dl;
d3, 13 each subequal to. ve; ratios of d4:14:d5:15, 3: 2i:iel; 15 twice the length of
al. ~ Ventral idiosoma: As in Pig. 27 B. “Legs: Setae dF, dG, dT of legs Iand iT
smooth; a' and a" IV each with 8 tines, a'l 2/3 length of a"I; sc3 and sc4 sub-
equal, extending beyond genua; vFI not ‘extending to ambulacrum:; vFIT extending
to ambulacrum; setae scl, sc2, aTIII, aTIV absent; tc'III and IV 1/2 length of
te"III and IV. Only the female is known.
Type material. From Pipilo erythrophthalmus (L.), Fringillidae: holotype
female, 1 female paratype, Lake Placid, Highlands County, Florida, July, 1960,
W.T. Atyeo, N. L. Braasch, K.R. Orwig. The holotype and 1 female paratype
are deposited with the University of Georgia, Athens, Georgia.
The name pipili refers to the host.
Syringophiloidus, new genus
Within the genera previously mentioned having MCA] parallel, Syringophi-
loidus is distinguished by the arrangement of the setae of the propodosomal region
and the loss of leg setae dFII and vs'II. This new genus appears most similar to
Niglarobia. These small to medium sized mites (630H- 875.) are found in the
secondaries of piciform birds of the family Picidae, and passeriform birds of the
following families: Bombycillidae, Cinclidae, Fringillidae, Icteridae, Laniidae,
Muscicapidae, Paridae, Parulidae, Ploceidae, and Turdidae. The name Syringo-
philoidus (masculine singular), meaning like Syringophilus, refers to the super-
ficial likeness to that genus.
Type species: Syringophilus minor Berlese, 1887, Acari, Myr. Scorp. a7,
10; ex. Passer domesticus (L.); Ploceidae, Passeriformes.
FEMALE. (1). Hypostomal apex smooth. (2). Lateral hypostomal teeth ab-
sent. (3). Chelicerae edentate. (4). Peritreme M-shaped; each lateral branch
with 2-4 chambers; each longitudinal branch with 6-14 chambers. (5). Stylophore
rounded posteriorly, usually extending below propodosomal plate. (6). Palpal
tibiotarsus rounded on distal margin. (7). Dorsal idiosomal setae smooth or
spinose; other setae smooth. (8). Propodosomal plate entire, rectangular in shape.
(9). Hysterosomal plate variable. (10). Setal pattern of propodosomal region
with six pairs of setae arranged 2-1-1-2. (11). Setae 12, 13, d3 long; subequal;
d3 closer to 12 than to13. (12). Setae d4, 14 long, subequal; d5, 15 short, sub-
equal. (13). ~ Genital series with two pairs ‘of setae; anal series variable, one or
two pairs of anal setae present. (14). Paragenital series with three pairs of
setae. (15). MCAlI parallel, not fused to MCA2. (16). Coxae III and IV moder-
ately sclerotized, crescent shaped. (17). Cuticular striations as in Fig. 28A, B;
transverse between coxae III and IV. (18). Legs I thicker than II; legs II sub-
equal in thickness to III and IV or legs I-IV subequal in thickness. (19). Leg
setae dGII, and vs'II absent. (20). Setae a' anda" multiserrate, 6-12 tines.
(21). Antaxial and id paraxial members of claw pair subequal; claws 1/3 length of
empodium. (22). Orders of hosts: Piciformes and Passeriformes. (23). Types of
Kethley: Syringophilidae
©)
Fig. 27. Aulonastus pipili new species. Female (holotype): A, dorsal
aspect; B, ventral aspect; C, idiosomal seta, smooth.
feathers inhabit: secondaries.
MALE. As in female except: (7). All setae smooth.
(10). Setal pattern of
propodosomal region 2-1-12 or 2-1-2-1 due to variable position Of 20. (th,
49
50 Contrib. AmersvEnt $1nsti vol. 5-)*no. 6, 1970
Setae 12, 13, d3 short. (12). Setae d4 short. (14). Paragenital series with two
pairs of setae; pgl absent; pg3 terminal. (15). MCAl weakly divergent. (17).
Cuticular striations as in Fig. 29A, B.
The genus Syringophiloidus includes 2 named and 18 new species. The named
species transferred to this genus are:
Syringophiloidus minor (Berlese)
new combination
(Figs. 28, 29)
Syringophilus minor Berlese, 1887, Acari, Myr. Scorp. 37, 10. - Fritsch, 1958,
Zool. Jahrbucher 86(3): 235, Fig. 5
Although all type material of this species is lost, material from the type host,
on loan from C.D. Radford and the Rocky Mountain Laboratory, was adjudged
conspecific and examined. Described originally from Passer domesticus (L.)
(Passeriformes: Plociedae), in Europe, this species has been collected from the
same host in England and Georgia.
Syringophiloidus seiurus (Clark)
new combination
Syringophilus seiuri Clark, 1964, Acarologia 6:81, Figs. 24, 25.
Although the holotype of this species is lost, 16 paratypes on loan from the
Rocky Mountain Laboratory were examined. Described originally from Seiurus
aurocapillus (L.) (Passeriformes: Parulidae), in Maryland, this species has been
collected from the same host in Florida and South Carolina, U.S.A.
Philoxanthornea, new genus
Among the genera having MCA] parallel to weakly divergent Philoxanthornea
is distinguished by the arrangement of the setae of the propodosomal region, the
loss of ieg setae dGII, vs'I and vs'IT, and the short length of 12, d3, 13. The
affinities of this new genus are uncertain. These small mites (520uU- 600) are
found in the flight feathers of charadriiform birds of the family Laridae and pele-
caniform birds of the family Phaethontidae. The name Philoxanthornea (feminine
singular) means lover of yellow birds, and refers to the name of the host birds.
Type species: Philoxanthornea anoa, new species; ex. Anous tenuirostris
(Temminck) 1823; Laridae, Charadriiformes.
FEMALE. (1). Hypostomal apex smooth. (2). Lateral hypostomal teeth absent.
(3). Chelicerae edentate. (4). Peritreme M-shaped; each lateral branch with 2-3
chambers; each longitudinal branch with 4-8 chambers. (5). Stylophore rounded
posteriorly; not extending to propodosomal plate. (6). Palpal tibiotarsus rounded
on distal margin. (7). All setae smooth. (8). Propodosomal plate weakly sclero-
tized margins indistinct, rectangular in shape. (9). Hysterosomal plate variable.
(10). Setal pattern of propodosomal region with six pairs of setae arranged 2-2-2.
(11). Setae 12, d3, 13 short; d3 closer to12 than to13. (12). Setae 14 long; d4,
d5, 15 short; d4, d5 on pygidial plate. (13). Genital series with two pairs of
setae; anal series with two pairs of setae. (14). Paragenital series with three
pairs of setae. (15). MCAl1 parallel to weakly divergent; fusion of MCA] and MCA2
indistinct. (16). Coxae III and IV weakly sclerotized, margins indistinct; artic-
ulations to trochanters prominent. (17). Cuticular striations as in Fig. 30A, B.
Kethley: Syringophilidae
Fig. 28. Syringophiloidus minor (Berlese). Female: A, dorsal aspect;
B, ventral aspect.
Fig. 29. Syringophiloidus minor (Berlese). Male: A, dorsal aspect;
B, ventral aspect.
Kethley: Syringophilidae 53
(18). Legs I thicker than II; legs II subequal in thickness to III and IV. (19). Leg
setae dGII, vs'I, vs'II absent. (20). Setae a' and a" multiserrate; 4-6 tines.
(21). Antaxial and paraxial members of claw pair subequal; with basal angle; claws
extending 1/3 length of empodium. (22). Orders of hosts: Charadriiformes and
Pelecaniformes. (23). Types of feathers inhabit: unknown.
MALE. As in female except: (9). Hysterosomal plate absent. (14). Para-
genital series with two pairs of setae, one pair absent. (15). MCA1 weakly di-
vergent. (21). Claws 1/4 length of empodium.
Philoxanthornea anoa, new species
(Fide .. Su Be)
FEMALE (holotype). Length 560u; width 145U. Gnathosoma: Hypostomal
apices unornamented, smooth. Peritremes, each lateral branch 1 chambered;
each longitudinal branch with 6-7 chambers. Dorsal idiosoma: Propodosomal
plate not divided, bearing setae vi, ve, sci, dl, 11; ratios of vi:ve:sci:sce,
1:2:41/2:5; dl, 11, sci subequal in length. Hysterosomal plate present, bearing
d3, 13, d4, d5; 12, d3, 13 each 11/2 times length of vi; ratios of d4:14:d5:15,
2:10:1:2/3; d4 three times length of vi. Ventral idiosoma: As in Fig. 30 B. Legs:
Setae dF, dG, dT of legs I and II smooth; a’ and a" I-IV with six tines each, al
1/2 length of a"I; sc3 and sc4 subequal, not extending beyond genua; vFII ex-
tending at least to ambulacrum; tc'III and IV subequal, each 1/2 length tc" III
and IV.
MALE (paratype). Length 450u; width 130uU. Gnathosoma: Hypostomal
apices unornamented. Peritremes as in female. Dorsal idiosoma: Propodosomal
plate not divided, bearing setae vi, ve, sci, dl, 11; ratios of vi:ve:sci:sce,
1:2:3:2; dl, 11, sci subequal in length. Hysterosomal plate absent; 12, d3, 13
subequal, 1/2 length of vi; ratios of d4:14:gl:g2, 3:18:1:1; d4 subequal to vi.
Ventral idiosoma and aedeagus: As in Fig. 31 B. Legs: As in female except a'l
subequal to a'll.
Type material. From Anous tenuirostris (Temminck), Laridae: Holotype fe-
male, 4 female paratypes, 2 male paratypes, Sand Island, Midway Atoll, Pacific
Ocean, February 20, 1963, collector unknown. Holotype and 2 female paratypes
and 1 male paratype deposited with U. S. National Museum, Washington D. C.;
remaining paratypes deposited with University of Georgia, Athens, Georgia.
Peristerophila, new genus
Species of Peristerophila may be distinguished from all other species of
Syringophilidae by the loss of setae vi, dFII, dFIII, dFIV, and vs'II, and the
shape of MCAl and MCA2. This new genus appears most similar to Aulonastus.
These small to medium sized mites (670U- 780) are found in the flight feathers
of columbiform birds of the family Columbidae. The name Peristerophila (feminine
singular) means pigeon lover, and refers to the host relationship of the genus.
Type species: Syringophilus columbae Hirst, 1920, Ann. Mag. Nat. Hist.
6:121-122; ex. Columba livia Gmelin; Columbidae, Columbiformes.
FEMALE. (1). Hypostomal apex ornamented; two pairs of median protuberances
present. (2). Lateral hypostomal teeth absent. (3). Chelicerae edentate. (4).
Peritreme M-shaped; each lateral branch with 2-3 chambers; each longitudinal
branch with 6-10 chambers. (5). Stylophore rounded posteriorly; not extending
below propodosomal plate. (6). Palpal tibiotarsus rounded on distal margin.
54 Contrib. Amer; Ent. Insts, vol..5,;no.6, 1970 -
Fig. 30. Philoxanthornea anoa new species. Female (holotype): A, dorsal
aspect; B, ventral aspect.
Kethley: Syringophilidae |
56 Contrib. Amer.-Enty inst. voles; 0. 6, 1970
(7). All setae smooth. (8). Propodosomal plate weakly sclerotized, margins indis-
tinct; entire, rectangular in shape. (9). Hysterosomal plate variable. (10). Setal
pattern of propodosomal region with five pairs (vi absent) arranged 1-1-1-2. (11).
Setae 12, 13, d3 long; d3 closer to 12 than to13. (12). Setae d4, 14 long, d31/2
to 3/4 length of 14; d4, 15 short; 14, 15 on common tubercle. (13). Genital series
with two pairs of setae; anal series with two pairs of setae. (14). Paragenital
series with three pairs of setae. (15). MCAlI parallel, not fused to MCA2. (16).
Coxae III and IV weakly sclerotized, margins indistinct. (17). Cuticular stria-
tions as in Fig. 32 A, B. (18). Legs I thicker than II; II subequal in thickness to
III and IV. (19). Leg setae dFII, dFIII, dFIV, and vs'Il absent. (20). Setae a'
and a" multiserrate, 16-24 tines. (21). Antaxial and paraxial members of claw
pair subequal; claws 1/2 length of empodium. (22). Order of hosts: Columbi-
formes. (23). Types of feathers inhabit: secondaries, primary coverts and sec-
ondary coverts.
MALE. As in female except: (1). Hypostomal apex unornamented. (10). Setal
pattern of propodosomal region 2-2-1; 11 not on propodosomal plate, position
variable. (11). Setae 13, d3 short. (12). Setae d4 short. (14). Paragenital series
with two pairs of setae, one pair of setae absent. (15). MCAl1 weakly divergent.
(17). Cuticular striations as in Fig. 33 A, B. (18). Legs I-IV subequal in thick-
ness. (21). Claws 1/3 length of empodium.
The genus Peristerophila includes two named and one new species. The
named species transferred to this genus are:
Peristerophila columba (Hirst)
new combination
Fig. 32)
Syringophilus columbae Hirst, 1920, Ann. Mag. Nat. Hist. 6:121-122; 1922,
Brit. Mus. (Nat. Hist.) Econ. Ser. 13: Fig. 41.
Although all type material of this species is lost, specimens on loan from
the University of Nebraska were adjudged conspecific and examined. Originally
described from Columbia livia Gmelin (Columbiformes: Columbidae) in Tennessee,
this species has been recollected in Kansas.
Peristerophila zenadoura (Clark)
new combination
Syringophilus zenadourae Clark, 1964, Acarologia, 6:83 Figs. 22, 23.
Although the holotype of this species is lost, one female paratype on loan
from the Rocky Mountain Laboratory was examined. Originally described from
Zenaidura macroura (L.) (Columbiformes: Columbidae) from Maryland, this species
has been recollected from the type host.
Syringonomus, new genus
Within the group of genera having the posterior portion of the stylophore con-
stricted, Syringonomus is distinguished by the annulate condition of the dorsal
idiosoma setae of the adult female, one pair of setae in the genital series and
the loss of leg setae vs'II. These small to medium sized mites (655y- 670) are
found in the flight feathers of procellariiform birds of the family Procellariidae
and charadriiform birds of the family Laridae. The name Syringonomus (masculine
Kethley: Syringophilidae | Sv
lOOu
AGS
Fig. 32. Peritserophila columba (Hirst). Female: A, dorsal aspect;
B, ventral aspect; C, a'IV.
58 Contrib. Amer. Ents Inst. vola 5 no. 6, .1970
eleyr
Fig. 33. Peristerophila sp. Male: A, dorsal aspect; B, ventral aspect.
singular) means inhabitant of tubes and refers to the avian host.
Type species: Syringqgnomus bulwerius, new species; ex. Bulweria bulweria
(Jardine and Selby); Procellariidae, Procellariiformes.
FEMALE. (1). Hypostomal apex slightly ornamented; four small median pro-
tuberances present. (2). Lateral hypostomal teeth absent. (3). Chelicerae eden-
tate. (4). Peritreme M-shaped; each lateral branch with 4-5 chambers; each lon-
gitudinal branch 14 chambered. (5). Stylophore constricted posteriorly, extending
beneath propodosomal plate. (6). Palpal tibiotarsus rounded on distal margin.
Kethley: Syringophilidae | 59
(7). Setae vi, ve, sci, sce, 11, dl, 12, 13, d3, AFI, dFII, dII annulate; other
setae smooth. (8). Propodosomal plate weakly sclerotized, margins indistinct.
(9). Hysterosomal plate absent. (10). Setal pattern of propodosomal region with
six pairs of setae arranged.2-2-2. (11). Setae 12, 13, d3 long, subequal; d3
closer to 12 than to13. (12). Setae d4, 14 long; d4 1/2 length of 14; d5, 15 short,
d4, d5 on pygidial plate. (13): Genital series with one pair of setae, one pair
of setae absent; anal series with two pairs of setae. (14). Paragenital series
with three pairs of setae. (15). MCAl weakly divergent, fused to MCA2. (16).
Coxae III and IV strongly sclerotized; articulation to trochanters prominent. (17).
Cuticular striations as in Fig. 34 A, B; longitudinal between coxae III and IV.
(18). Legs I thicker than II; legs II thicker than III and IV. (19). Leg setae vs'II
absent. (20). Setae a' and a" multiserrate, 6-8 tines. (21). Antaxial and par-
axial members of claw pair subequal, with basal angle; claws 1/2 length of
empodium. (22). Orders of hosts: Procellariiformes and Charadriiformes. (23).
Types of feathers inhabit: unknown.
MALE. As in female except: (5). Stylophore not extending under propodoso-
mal plate. (7). All setae smooth. (10). Setal pattern of propodosomal region 3-
2-1. (11). Setae 13, d3 short. (12). Setae d4 short. (14). Setae pg3 terminal.
(16). Coxae III and IV weakly sclerotized, margins indistinct. (17). Cuticular
striations as in Fig. 35 A. B. (21). Claws 1/3 length of empodium.
The genus Syringonomus includes one new species:
Syringonomus bulwerius, new species
Fics, 24,35)
FEMALE (holotype). Length 690uU; width 1904. Gnathosoma: Hypostomal
apices slightly ornamented, two pairs of short protuberances present. Peritremes,
each lateral branch 4 chambered; each longitudinal branch 12 chambered. Dorsal
idiosoma: Propodosomal plate not divided, bearing setae vi, ve, sci, dl, 11;
ratios of vi:ve:sci:sce, 1:1:1:2; dl, 11, sce subequal in length. Hysterosomal
plate present, bearing d3, 13; 12, 13 subequal; d3 3/4 length of 12; ratios of d4:
14:05:15, 9:15:1:1; d4 13/4 times length of dl. Ventral idiosoma: As in Fig. 34B.
Legs: Setae dF, dG, AT of legs Iand II annulate; a’ and a" I-IV with eight tines
each, a'l 2/31 length of | a"l; sc3 and sc4 subequal, not extending beyond genua;
vFIT extending at least to ambulacrum; | tc'III and IV 2/3 length of tc"III and IV.
MALE (paratype). Length 530; ‘width 180. Gnathosoma: Hypostomal
apices unornamented. Peritremes as in female. Dorsal idiosoma: Propodosomal
plate not divided, bearing setae vi, ve, dl; ratios of vi:ve:sci:sce, 1:2:2:2; 11
11/2 times length of vi; dl 11/4 times length of vi. Hysterosomal plate absent;
a3, 13 subequal to length of vi; 12 11/2 length of vi; ratios of d4:l4:gl:g2, 5:30:
1:1; d4 twice length of vi. Ventral idiosoma and aedeagus: As in Fig. 35 B.
Legs: As in female except setae dF, dG, dT of legs I and II smooth; a'I subequal
to a'll,
Type material. From Bulweria bulweria (Jardine and Selby), Procellariidae:
holotype female, 2 male, 10 female paratypes, Nihoa Island, Hawaiian Islands,
June 6, 1963, collector unknown. Holotype, 1 male and 5 female paratypes de-
posited with U.S. National Museum, Washington, D.C., additional paratypes
deposited with University of Georgia, Athens, Georgia.
The name bulwerius refers to the host.
60 Contrib. Amer.:Ents Inst... vol<.5;-ho. 6, 1970
l\OOu
Fig. 34. Syringonomus bulwerius new species. Female (holotype): A,
dorsal aspect; B, ventral aspect; C, idiosomal seta, annulate.
Kethely: Syringophilidae
Fig. 35. Syringonomus bulwerius new species.
dorsal aspect; B, ventral aspect.
Male (paratype): A,
61
62 Contrib. Amer. Ent.) Inst., vol. 5,:no..6, 1970
Stibarokris, new genus
Among the previously mentioned genera having a constriction of the posterior
margin of the stylophore, Stibarokris is distinguished by the presence of lateral
hypostomal teeth, and the absence of leg setae vs'II. Although this new genus
is grouped with those genera having a common stylophore configuration, the pres-
ence of the lateral hypostomal teeth is a unique feature among the Syringophilidae.
These medium sized mites (650U- 790) are found in the covert feathers of
pelecaniform birds of the family Phalacrocoracidae and ciconiiform birds of the
family Phoenicopteridae. The name Stibarokris (masculine singular) means sturdy
projection and refers to the lateral hypostomal teeth.
Type species: Stibarokris phalacrus, new species, ex. Phalacrocorax auritus
(Lesson), 1831; Phalacrocoracidae, Pelecaniformes.
FEMALE. (1). Hypostomal apex unornamented. (2). Lateral hypostomal teeth
present. (3). Chelicerae edentate. (4). Peritreme M-shaped; each lateral branch
with 6-7 chambers; each longitudinal branch with 10-16 chambers. (5). Stylophore
constricted posteriorly, extending beneath propodosomal plate. (6). Palpal tibio-
setae smooth. (8). Propodosomal plate entire, rectangular in shape. (9). Hystero-
somal plate bears 12, d3. (10). Setal pattern of propodosomal region with six
pairs of setae arranged 2-2-2. (11). Setae 12, d3, 13 long; d3 closer to 12 than
to 13. (12). Setae d4, 14 long, d4 1/2 length of 14; d5, 15 short. (13). Genital
series with two pairs of setae; anal series with two pairs of setae. (14). Para-
genital series with three pairs of setae. (15). MCA1 weakly divergent, fused to
MCA2 indistinctly. (16). Coxae III and IV strongly sclerotized, rectangular.
(17). Cuticular striations as in Fig. 36 A, B. (18). Legs I thicker than legs II;
legs II thicker than III and IV. (19). Leg setae vs'II absent. (20). Setae a' and
a" multiserrate, 8-14 tines. (21). Antaxial and paraxial members of claw pair
subequal; claws 1/2 length of empodium. (22). Orders of hosts: Pelecaniformes
and Ciconiiformes. (23). Types of feathers inhabit: coverts.
MALE. Unknown.
Stibarokris phalacrus, new species
(Fig. 36)
FEMALE (holotype). Length 800uU; width 245u. Gnathosoma: Hypostomal
apices unornamented, smooth (Fig. 36 C, D). Peritremes, each lateral branch
with 5-6 chambers; each longitudinal branch 12 chambered. Dorsal idiosoma:
Propodosomal plate not divided bearing setae vi, ve, sci, dl, 1]; ratios of vi:
ve:sci:sce, 1:11/8:11/4:1 1/2; 11 subequal to sce; dl 11/6 lengthll. Hystero-
somal plate divided, restricted to bases of d3; 12, d3, 13, subequal to sce; ratios
of d4:14:d5:15, 8:32:1:2; d4 subequal to length of 11. Ventral idiosoma: As in
Fig. 36 B. Legs: Setae dF, dG, dT of legs I and II smooth; a' and a" I-IV with
8 tines each, a'I subequal to a"I; sc3 extending beyond genu III; sc4 not extend-
ing beyond genu IV; vFI not extending to ambulacrum; vFII extending at least to
ambulacrum; tc'III and IV 1/2 length tc"III and IV. Only the female is known.
Type material. From Phalacrocorax auritus (Lesson), Phalacrocoracidae:
holotype female, 7 female paratypes, Pigeon Key, Monroe County, Florida,
November 28, 1967, R. Heard, W. B. Sikora.
The name phalacrus refers to the host.
Kethley: Syringophilidae 63
Fig. 36. Stibarokris phalacrus new species. Female (holotype): A,
dorsal aspect; B, ventral aspect; C, hypostomal lips, dorsal aspect;
D, hypostomal lips, ventral aspect; E, idiosomal seta, knobbed.
64 Contrib. Amer, Ent: Inst... vol.-5;-no. 6, 1970
Genus Picobia Haller
Picobia Haller, 1878, Zeit. wiss. Zool. 30:81-91. - Oudemans, 1906, Mem. Soc.
Zool, Paris, 19:73.-~ Dubinin, 1957, Mag. Parasitol. Inst; Zool. Akad. Sc.
USSR. 5 17271-1536:
The truncate condition of the palpal tibiotarsus, thickened legs I and II, and
the loss of leg setae scl, sc2, dFIII and dFIV distinguishes Picobia from all other
genera of Syringophilidae. This genus is strikingly different from all other sy-
ringophilid genera and its affinities are obscured by host adaptations and con-
comitant structural modifications. These small to medium sized mites (610u-
780u.) are found subcutaneously in the epidermal wing tissue of piciform birds of
the family Picidae (Haller 1878), and have been taken from birds in the following
families: Apodidae, Apodiformes; Phasianidae, Galliformes; Columbidae, Columbi-
formes; Sturnidae and Turdidae, Passerifromes; and Upupidae, Coraciiformes.
Type Species: Picobia heeri Haller, 1878, ex. Picus canus Gmelin (=Gecinus
canus), Picidae, Piciformes. (Type by original designation.)
FEMALE. (1). Hypostomal apex unornamented. (2). Lateral hypostomal teeth
absent. (3). Chelicerae edentate. (4). Peritreme M-shaped; each lateral branch
with 2-6 chambers; each longitudinal branch with 4-20 chambers. (5). Stylophore
elongate, rounded posteriorly; variable relation to propodosomal plate. (6). Palpal
tibiotarsus truncate (Fig. 2 B), segmental sutures indistinct. (7). Setae variable,
spinose, knobbed, annulate, or smooth. (8). Propodosomal plate variable, di-
vided, entire; sclerotization usually restricted to bases of vi, ve, sci. (9). Hy-
sterosomal plate variable. (10). Setal pattern of propodosomal region with six
pairs of setae arranged 2-2-2. (11). Setae 12, d3, 13 long; d3 closer to 12 or
equidistant between 12 and 13. (12). Setae 14 long; d4, d5, 15 short. (13). Gen-
ital series with two pairs of setae; anal series with one pair of setae. (14). Para-
genital series with three pairs of setae. (15). MCAI variable, parallel, weakly
convergent, or weakly divergent; MCA] not fused to MCA2; MCA2 produced pos-
teriorly beyond coxal field. (16). Coxae III and IV strongly sclerotized; posterior
margin strong; articulation with trochanters prominent. (17). Cuticular striations
as in Fig. 37 A, B; transverse between setal bases 3a; transverse between coxae
ITI-IV. (18) . Legs I and II subequal in thickness, 2-3 times thicker than III and
IV. (19). Leg setae scl, sc2, GFIII and dFIV absent. (20). Setae a' and a" var-
iable, rodlike on legs III-IV or absent; rodlike or irregularly serrate with 1-2
points on legs I-II. (21). Antaxial and paraxial members of claw pair III-IV var-
iable, subequal or dissimilar; claws extending 1/4 length of empodium. (22).
Orders of hosts: Apodiformes, Columbiformes, Coraciiformes, Galliformes,
Passeriformes, Piciformes, and Podicipediformes. (23). Types of feathers in-
habit: unknown; P. heeri subcutaneous in wing tissue of Picus canus.
MALE.. As in female except: (7). All setae smooth. (11). Setae d3, 13 short;
d3 closer to 13 than to12. (14). Paragenital series with two pairs of setae.
The genus Picobia includes 3 named and 6 new species. The named species
are:
Picobia heeri Haller
Only the pertinent citations of this species are given.
Picobia heeri Haller, 1878, Zeit. wiss. Zool. 30:81-91. - Oudemans, 1906, Mem.
Soc. Zool. Paris, 19:73. - Dubinin, 1957, Mag. Parasitol. Inst. Zool. Akad.
Sc. UseR., 17: 71=1S6
Kethley: Syringophilidae | 65
All type material of this species originally collected in Germany is lost, and.
representative material was not available for study.
Picobia dryobatis (Fritsch)
new combination
Syringophilus dryobatis Fritsch, 1958, Zool. Jahrbucher 86 (3):242-243, Fig. 10.
| All the type material of this species originally collected from Dendrocopus
major (L.), Picidae in Germany is lost, and representative specimens were not
available for study.
Picobia zumpti (Lawrence)
new combination
(Fig. 37)
Syringophilus zumpti Lawrence, 1959, Parasitology, 49:425, Fig. 5.
The holotype of this species, on loan from the Natal Museum, was examined.
Originally described from Streptopelia capicola (Sundemall) (Columbiformes:
Columbidae) in South Africa, this species has not been recollected.
Owing to the paucity of the descritpions, the following species cannot be
assigned with certainty to any of the existing genera:
Syringophilus anthi Fritsch, 1958, Zool. Jahrbucher 86(3):240-242, Fig. 9. ex.
Anthus trivialis (L.) (Passeriformes: Motacillidae), from Germany.
Syringophilus bisetatus Fritsch, 1958, Zool. Jahrbucher 86(3):238, Fig. 7. ex.
Sylvia communis Latham (Passeriformes: Sylviidae), from Germany.
Syringophilus glandarii Fritsch, 1958, Zool. Jahrbucher 86(3):235-238, Fig. 6.
ex. Garrulus glandarius (L.) (Passeriformes: Corvidae), from Germany.
Ssyringophilus major Berlese, 1878, Ac. Myr. Scorp. Ital., fasc. 37, no. 9 and
10. ex. Fringilla montifringilla (L.) (Passeriformes: Fringillidae), from Europe.
Syringophilus troglodytis Fritsch, 1958, Zool. Jahrbucher 86(3):238-240, Fig. 8.
ex. Troglodytes troglodytes (L.) (Passeriformes: Troglodytidae), from Germany.
DEVELOPMENT
The life history stages are typical of the cheyletoids (Hughes, 1961), com-
prising the egg, larva, and two nymphal stages prior to the adult. The deuto-
nymphal stage is lacking. Although genital setae are still lacking, leg chaeto-
taxy is complete in the tritonymph, except for sc4.
A complete series of the life stages was not available for all the species
studied. There are differences in the pattern of addition of setae in the immature
stages. However, the following conditions hold for all genera: dGI is added in
the protonymph; vs'I, vs'II, vs'III and dFII are either present or absent in the
larva--if absent they are not added; dGII, if present in the adult, is added in the
protonymph.
The developmental pattern of Syringophilus bipectinatus Heller, 1880,
66 Contrib. Amer. Ent. Inst., vol. 5, no. 6, 1970
oA
m
ak
Fig. 37. Picobia zumpti (Lawrence). Female (holotype): A, dorsal aspect;
B, ventral aspect; C, claws tarsus IV; D, a'IIl; E, a'l.
(Table II), was chosen for illustrative purposes because in this species most of
the diagnostic characters employed in generic classification are present. In
addition, since S. bipectinatus is the type species for the family, it is employed
here as an arbitrary standard to which other members of the family may be com-
pared.
67
Kethley: Syringophilidae
A, dorsal aspect; B, ventral aspect.
Fig. 38. Picobia sp. Male:
68 Contrib. Amer. Ent. Insts, vol. 5, no. 6, 1970
TABLE II
Leg chaetotaxy of developmental stages of
Syringophilus bipectinatus Heller
The following setae are present in the larva:
Coxae 1b, 2a, 3a.
Trochanters none
Femora @bi-itit,-yrl-I1.
Genua 1GI-II, 1G"III, sigma
Tibiae Gilli, vil-til, 1 I-H, 1?" i-10,. pai.
Tarsi Ui-ti ou i-1l ve li, ft, te il, to Pol, Pe’, PB", omega i-il.
The following setae are added in the protonymph:
Coxae ic, 20; 3b; 30:
Trochanters none
Femora OFIV.
Genua aGli-il;
Tibiae QTV, vil¥, Ii" Iv.
Tarsi TC) 1C"l, a'l-Ill, a" ell, ulv, av, te'lV, 2c!
The following setae are added in the tritonymph:
Coxae 4b, 4c.
Trochangers §¢l, scc, sc3.
Femora none
Genua iG Ty
Tibiae none
Tarsi aiv, a iV.
The following seta is added in the adult:
Trochanters sc4.
NOTE: The eupathid pair (TC) changes from setiform in the larva (tc) to true
eupathid type in the protonymph; (P) is eupathid on the larva.
Kethley: Syringophilidae — 69
BIONOMICS AND HOST-PARASITE RELATIONS
All known species of syringophilids are found inside the calmus of wing
feathers (primaries, secondaries, and coverts), or in the case of Picobia heeri,
in the subcutaneous tissue of the wing of Picus canus.
The mites remain inside the calmus, and leave only at the onset of molting
to seek new feathers. Thus, reproduction and development take place within the
quill. All life stages feed by inserting the long, needle-like chelicerae through
the fibrous wall of the calmus into the flesh of the wing. The haplodiploid sys-
tem is used to limit the number of males to one for every ten to fifteen females.
Dispersal from an old, about to be molted, feather to a newly developing feather
is accomplished by females presumed to be non-gravid , but fertilized. Either
the mites enter the new calmus when the superior umbilicus of the developing
feather is unobstructed or they make an entrance hole in the region of the supe-
rior umbilicus.
During dispersal, the mites exhibit a preference for certain feathers. The
phenomenon of site selection may be explained in terms of two components--a
feather component and a mite component--each involving several variables.
The feather varies in volume of the calmus; calmus wall thickness; fre-
quency of feather milts per breeding cycle of host; pattern or sequence of molt-
ing of the feathers within the various tracts and the coordination between tracts.
The mites vary as to: volume of a female mite; fecundity; volume occupied by
One generation of progeny; number of generations per quill molt; number of females
invading a quill; reproductive interaction among females in instances of multiple
invasions; mobility of the mites, and the ability to create a hole in the calmus
for entry. Combinations of the above factors appear to limit the mites to specific
feathers and offer the basis for the evolution of feather preferences among the
mites.
Indeed, adaptations restrict mites to specific conditions so that some genera
are limited to specific feather tracts. For example, an individual passerine bird
may have a species of Syringophilopsis in the primaries, a species of Syringo-
philoidus in the secondaries, and a species of Aulonastus in the coverts. Simul-
taneous collection of species representing two genera of quill mites from the
same individual host has occured from Turdus migratorius, Agelaius phoeniceus,
Passer domesticus, and Sturnus vulgaris. In addition, representatives of two
genera are known from the same species of host for five other species of syringo-
philids, although the mites were not collected from the same host individual.
Molting by the host forces mites to move from quill to quill. In contrast, the
movement of mites from host to host is governed by the periods and types of con-
tact between hosts. Thus, the events of mating and nesting, and periods of gre-
gariousness of the host afford opportunities of possible transmission. Most of
the transfers between individual birds may occur during the nesting period of the
host. The timing of these transfers must be correlated with development of the
offspring at hatching, and the length of exposure of the immature birdto the adults.
The genera of Syringophilidae are now known to parasitize sixteen orders of
birds. For the most part, the genera appear to be specific at the ordinal level of
the avian host. Of the seven cases of ordinal overlap, only one lacks a possible
explanation. The ordinal crossing-over can be explained in the same way used
by K. C. Emerson (1962), and T. Clay (1949,.1950, 1957) to resolve confusing
host associations of bird lice.
Of the one hundred twenty-two species of quill mites examined, only eight
are known from more than one host species. Examination of additional hosts may
70 Contrib. Amer. Ent. Inst., vol. 5, no. 6, 1970
show that this initial sampling suggests too great a host specificity. It is ap-
parent that the current study represents only a beginning, and that an enormous
fauna is unknown.
Kethley: Syringophilidae
HOST-PARASITE LIST
ORDER: PROCELLARIIFORMES
Bulweria bulweria (Jardine and Selby), 1828
Syringonomus bulwerius new species
ORDER: PELECANIFORMES
Phalacrocorax auritus (Lesson), 1831
Stibarokris phalacrus new species
ORDER: CICONIIFORMES
Casmerodius albus egretta (Ord), 1825
Trypetoptila casmerodia new species
ORDER: ANSERIFORMES
Branta canadensis (L.), 1758
Chenophila branta new species
ORDER: GALLIFORMES
Gallus gallus (L.)
Syringophilus bipectinatus Heller
ORDER: GRUIFORMES
Gallinula chloropus (L.), 1758
Ascetomylla gallinula new species
ORDER: CHARADRIIFORMES
Charadrius wilsonia wilsonia Ord, 1814
Selenonycha baltoda new species
Aramus scolopaceus (Gmelin), 1789
Niglarobia trouessarti (Oudemans)
Ereunetes pusillus (L.), 1766
Niglarobia ereuneti new species
Tringa ochropus (L.) 1758
Niglarobia helleri (Oudemans)
Tringa totanus (L.), 1758
Creagonycha totana (Oudemans)
Anous tenuirostris (Temminck)
Philoxanthornea anoa new species
Larus delawarensis Ord, 1815
Creagonycha lara new species
ORDER: COLUMBIFORMES
Columbia livia Gmelin, 1789
Peristerophila columbae (Hirst)
Streptopelia capicola (Sundemall), 1857
Picobia zumpti (Lawrence)
Zenaidura macroura (L.), 1758
Peristerophila zenadoura (Clark)
71
Procellariidae
Phalacrocoracidae
Ardeidae
Anatidae
Phasianidae
Rallidae
Charadriidae
Scolopacidae
Laridae
Columbidae
72 Contrib. Amer. Ent..Inst., vol..5, mo. 6; 1970
ORDER? PICIFORMES
Dendrocopus major (L.), 1758
Picobia dryobatis (Fritsch)
Picus canus Gmelin, 1788
Picobia heeri Haller
ORDER: PASSERIFORMES
Mimus polyglottos (L.), 1758
Torotrogla mima new species
Hylocichla fuscescens (Stephans), 1817
Syringophilopsis hylocichla (Clark)
Turdus pilaris (L.), 1758
Syringophilopsis turdus (Fritsch)
Phainopepla nitens (Swainson), 1837
Torotrogla villosa (Hancock)
Dendroica coronata (L.), 1766
Aulobia dendroicus (Clark)
Seiurus aurocapillus (L.), 1766
Syringophiloidus seiurus (Clark)
Passer domesticus (L.), 1758
Syringophiloidus minor (Berlese)
Agelaius phoeniceus (L.), 1766
Syringophilopsis elongatus (Ewing)
Carduelis carduelis (L.), 1758
Syringophilopsis fringilla (Fritsch)
Fringilla coelebs (L.), 1758
Syringophilopsis fringilla (Fritsch)
Passerina cyanea (L.), 1766
Syringophilopsis passerina (Clark)
Pipilo erythrophthalmus (L.), 1758
Aulonastus pipili new species
Picidae
Mimidae
Turdidae ©
Ptilgonatidae
Parulidae
Ploceidae
Icteridae
Fringillidae
Kethley: Syringophilidae vs
There is considerable literature concerned with the diagnosis and treatment
of domestic avian parasites. Comments concerning Syringophilus bipectinatus,
which occurs on domestic chickens, and P. columbae from pigeons, have ap-
peared in veterinary publications and are extremely numerous that only a few are
given below.
Baker, E. W. 1949. A review of the mites of the family Cheyletidae in the United
States National Museum. Proc. U.S. Natl. Mus. 99(3238): 267-320.
Baker, E. W. 1965. A review of the genera of the family Tydeidae (Acarina).
Advances in Acarology II, Cornell University Press Ithaca, New York.
Baker, E. W. and G. W. Wharton. 1952. An introduction to acarology. Mac-
millan Co., pp. 146-258
Baker, .E. Wi, Te M. Evans, De JicGould) W. BY Hull2-and He 1: Keegan. 1956.
A manual of parasitic mites of medical or economic importance. New York
156-158.
Baker, Es Wij J<H.:Camin,:Po-Cunliife;, 7. Ay: Woolley, CLE. Yunker. Guide
to the families of mites. Contribution no. 3, Institute of Acarology,
College Park, Md. 1958 1-242,
Beer, R. E. 1954. A revision of the Tarsonemidae of the western hemisphere
(Order Acarina). Univ. Kansas Sci. Bull. 36: 1091-1387.
Benbrook, E. A. 1955. Veterinary Clinical Parasitology. Iowa State College
Press, Ames, Iowa. p. 116.
Berlese, A. 1887. Acari, Myriopoda et Scorpiones, Padova. fasc. 377 ROS.
Be Oy
Berlese, A. 1893. Acari, Myriopoda et Scorpiones, Ordo Prostigmata. Portici.
pe 161%
Camin, J. H., et. al. 1967. A new family of cheyletoid mites from the cloaca
of aquatic turtles (Acari: Acariformes: Eleutherengona). J. Med. Entomol.
4(3): 261-272.
Clark, G. M. 1964. The acarine genus Syringophilus in North American birds.
Acarologia 6: 77-92.
Clay, T. A. 1949. Some problems in the evolution of a group of ectoparasites.
Evolution 3: 279-299.
Clay, T. A. 1950. A preliminary survey of the distribution of the Mallophaga
(Feather lice) on the class Aves (birds). Bombay Nat. Hist. Soc. 49:
429-443.
Clay, T., 1957. in International Symposium on Host specificity among parasites
of vertebrates. lst, Universite do Neuchatel. pp. 120-156.
Cunliffe, F. 1955. A proposed classification of the trombidiforme mites (Acarina).
Proc. Entomol. Soc. Washington. 57: 209-218.
Dubinin, W. B. 1957. A new classification of the mites of the superfamilies
Cheyletoidea W. Dub. and Demodicoidea W. Dub. (Acariformes, Trombedi-
formes). Akad. nauk S.S.S.R., Zool. Inst., Parazitol. Sborn., 17: 17-136.
Ewing, H. E. 1911. Predaceous and parasitic Acarina. Psyche, 18: 37-43.
Ewing, H. E. 1938. North American mites of the subfamily Myobiinae, new sub-
family (Arachnida). Proc. Entomol. Soc. Washington 40: 180-197.
Fritsch, W. 1958. Die Milbengattung Syringophilus Heller 1880 (Subordo Trom-
bidiformes, Fam. Myobiidae Megnin 877). Zool. Jahrbucher 86: 227-244.
Grandjean, F. 1944. Observations sur les Acariens de la famille des Stigmaeidae.
Arch. Sci. phys. nat. 26: 103-131.
Haller, G. 1877. Zeit. wiss. Zool. (Siebold und Kolliker: Leipzig), 30: 91-93.
Hancock, J. L. 1895. Anew Trombidian. Amer. Nat. 29: 382-385.
74 Contrib. Ameri Ents Inst. , volu 5,hno..6, 1970
Hancock, J. L. 1895. Picobia villosa (Hancock). Amer. Nat. 29: 866-867.
Heller, A. 1880. Die Schmarotzer, Munich und Leipzig p. 186-188.
Hering, M. 1934. Syringophilus lhommi, in Tenthredinoides, L'Amateur do
Papillons 7: 154-160.
Hirst, S. 1920. Notes on Acari parasitic on birds with descriptions of two new
species. Ann. Mag. Nat. Hist. 6: 121-122.
Hirst, S. 1922. Mites injurious to domestic animals. Brit. Mus. (Nat. Hist.)
Econ. Ser. 13: 1-107.
Hughes, A. M. 1961. The Mites of Stored Food. Min. Agric. Fish., Food,.
Tech. Bull., No. 9, p. 178.
Lapage, G. 1956. Veterinary Parasitology, Oliver and Boyd, Edingurugh and
London, p. 683.
Lavoipierre, M. M. J. 1953. The undescribed male and female of the pigeon
quill mite, Syringophilus columbae, Hirst 1920. Trans. Roy. Soc. Trop.
Med. Hygiene 47: 7. ;
Lawrence, R. F. 1959. New mite parasites of African Birds. Parasitology 49:
425-427.
Lipovsky, L. J. 1953. Polyvinyl alcohol with lacto-phenol, a mounting and
clearing medium for chigger mites. Ent. News 64: 42-44.
Oudemans, A. C. 1904a. Acarologische Aanteekenigen, XIII. Entomol. Ber.,
1(19): 171.
Oudemans, A. C. 1904b. Acarologische Aanteekenigen, XIV. Entomol. Ber.,
1(20): 190-191.
Oudemans, A. C. 1905. Acarologische Aanteekenigen, XV. Entomol. Ber.,
P12 TS 207%
Oudemans, A. C. 1906. Revision des Cheyletines. Mem. Soc. Zool. 19: 36-144.
Peters, J. L. 1931-1960. Check-list of birds of the world. Vol. I-VII, IX, XV.
Harbard Univ. Press, Cambridge.
Wetmore, A. 1906. A classification for the birds of the world, Smithsonian
Misc. Coll., 139: 1-37.
albus, Casmerodius 25, 28, 71
ALUDIDAE 18
ANATIDAE 37, 71
ANSERIFORMES 37, 71
APODIDAE 64
APODIFORMES 64
ARDEIDAE 25, 71
auritus, Phalacrocorax 62, 71
aurocapillus, Seiurus 50, 72
BO MBYCILLIDAE 48
BUCEROTIDAE 24
bulweria, Bulweria 58, 59, 71
canadensis, Branta 37, 39, 71
canus, Picus 64, 69, 72
capicola, Streptopelia 65, 71
carduelis, Carduelis 21, 72
CHARADRIIDAE 31, 71
CHARADRIIFORMES 33, 35, 44, 53, 59, 71
chloropus, Gallinula 24, 25, 71
CICONIIFORMES 25, 62, 71
CINCLIDAE 48
coelebs, Fringilla 21, 72
COLUMBIDAE 53, 64, 71
COLUMBIFORMES 56, 64, 71
communis Sylvia 65
CORACTIFORMES 64
coronata, Dendroica 41, 72
cyanea, Passerina 23, 72
delawarensis, Larus 34, 37, 71
domesticus, Passer 48, 50, 69, 72
erythrophthalmus, Pipilo 47, 48, 72
FRINGILLIDAE 18, 28, 47, 48, 72
fuscescens, Hylocichla 21, 72
GALLIFORMES 17, 18, 64, 71
gallus, Gallus 17, 18, 71
glandarius, Garrulus 65
GRUIFORMES 24, 71
HIRUNDINIDAE 18
ICTERIDAE 18, 48, 72
LANIIDAE 48
LABIA, 31, 34, 60, 58,71
livia, Columba 53, 56, 71
macroura, Zenaidura 56, 71
major, Dendrocopus 65, 72
migratorius, Turdus 69
MIMIDAE 28, 72
montifringilla, Fringilla 65
MUSCICAPIDAE 48
nitens, Phainopepla 30, 72
Numididae 17
ochropus, Tringa 47, 71
PARIDAE 48
PARULIDAE 18, 41, 48, 72
PASSERIFORMES 21, 29, 41, 47, 48, 64, 72
PELECANIFORMES 53, 62, 71
HOST INDEX
PHAETHONTIDAE 50
PHALACROCORACIDAE 62, 71
PHASIANIDAE 17, 18, 64, 71
phoeniceus, Agelaius 21, 69, 72
PHOENICOPTERIDAE 62
PICIDAE 47, 48, 72
PICIFORMES 47, 48, 72
pilaris, Turdus 24, 72
PLOCEIDAE 48, 72
polyglottos, Mimus 28, 31, 72
PROCELLARIIDAE 56, 71
PROCELLARIIFORMES 59, 71
PTILGONATIDAE 28, 72
pusillus, Ereunetes 44, 71
PYCNONOTIDAE 18
RALLIDAE 24, 71
scolopaceus, Aramus 47
SCOLOPACIDAE 31, 34, 43, 71
STURNIDAE 64
SYLVIIDAE 41
tenuirostris, Anous 50, 53, 71
totanus, Tringa 37, 71
trivialis, Anthus 65
troglodytes, Troglodytes 65
TURDIDAE 18, 28, 48, 64, 72
TYRANNIDAE 18
UPUPIDAE 64
VIREONIDAE 18
vulgaris, Sturnus 69
wilsonia, Charadrius 31, 34, 71
75
76
PARASITE INDEX
In the following index, all names are listed alphabetically, regardless of whether they are
genus-group or species-group names. An asterisk * indicates a synonym. The underlined pages
are those which begin the primary treatment of the genus or species. Numbers in parentheses refer
to the figures illustrating the species in question.
anoa (Philoxanthornea) 50, 53, 71, (30, 31)
anthi (Syringophilus) 65
Ascetomylla 16, 17, 24
Aulobia 17, 41, 43
Aulonastus 16, 41, 47, 53, 69
baltoda (Selenonycha) 12, 31, 34, 71, (18, 19)
bipectinatus (Syringophilus) 12, 14, 17, 18-20,
65-67, 71, 73, (8; 9)
bisetatus (Syringophilus) 65
branta (Chenophila) 37, 38, 71, (22)
bulwerius (Syringonomus) 58, 59, 71, (34, 35)
casmerodia (Trypetoptila) 25, 28, 71, (14, 15)
Chenophila 16, 37
columba (Peristerophila) 56, 71, 73, (32)
*columbae (Syringophilus) 53, 56
Creagonycha 17, 28, 31, 34
*dendroicae (Syringophilus) 41
dendroicus (Aulobia) 7, 41, 72, (23, 24)
dryobatis (Picobia) 65, 72
*dryobatis (Syringophilus) 65
elongatus (Syringophilopsis) 12, 21, 22, 23,
72, 00, 11)
*elongatus (Syringophilus) 18, 21
ereuneti (Niglarobia) 44, 71, (25, 26)
fringilla (Syringophilopsis) 21, 72
*fringillae (Syringophilus) 21
gallinula (Ascetomylla) 24, 25, 71, (12, 13)
glandarii (Syringophilus) 65
Harpypalpus 6
Harpyrhynchus 6
heeri (Picobia 64, 69, 72
helleri (Niglarobia) 47, 71
*helleri (Syringophilus) 47
hylocichla (Syringophilopsis) 21, 72
*hylocichlae (Syringophilus) 21
*icteridae (Syringophilus) 21
lara (Creagonycha) 34, 36, 71, (20, 21)
major, (Syringophilus) 65
mima (Torotrogla) 28, 31, 72, (16, 17)
minor (Syringophiloidus) 12, 50, 72, (28, 29)
*minor (Syringophilus) 48, 50
Niglarobia 16, 41, 43, 48
passerina (Syringophilopsis) 23, 72
*passerinae (Syringophilus) 23
Peristerophila 16, 53
phalacrus (Stibarokris) 62, 71, (36)
Philoxanthornea 16, 41, 50
Picobia 4,5,16,64
pipili (Aulonastus) 47, 48, 72, (27)
seiurus (Syringophiloidus) 50, 72
*seiuri (Syringophilus) 50
Selenonycha 17, 28, 31, 34
Stibarokris 16, 37, 62
Syringonomus 16, 37, 56
Syringophilidae 6
Syringophiloidus 16, 41, 48, 69
Syringephilopsis:17,:18,24,°25,° 69
Syringophilus 4,5, 17, 48
Torotrogla 16, 28
totana (Creagonycha) 37, 71
*totani (Syringophilus) 37
troglodytis (Syringophilus) 65
trouessarti (Niglarobia) 47, 71
*trouessarti (Syringophilus) 47
Trypetoptila 16, 17, 25
*turdi (Syringophilus) 24
turdus (Syringophilopsis) 24, 72
*villosa (Picobia) 30
villosa (Torotrogla) 30, 72
zenadoura (Peristerophila) 56, 71
*zenadourae (Syringophilus) 56
zumpti (Picobia) 8, 65, 72, (2B, 37)
*zumpti (Syringophilus) 65
(Continued from inside front cover)
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CONTRIBUTIONS TO THE MOSQUITO FAUNA OF
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The genus Hezzmannia Ludlow in Southeast Asia
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P. F. Mattingly
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Contributions
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Volume 5, Number 7, 1970
CONTRIBUTIONS TO THE MOSQUITO FAUNA OF
SOUTHEAST ASIA. - VI
The genus Heizmannia Ludlow in Southeast Asia
by
P. F. Mattingly
CONTENTS *
PL PE EI IN ei ys ura tee ST eee EP a gE get he inh ae SS
GENUS HEIZMANNIA . . . BO igs ee ta ean We gig at Gh lage a Mag agth nee
KEYS TO SOUTHEAST ASIAN SPECIES
eS ee ee a he a ea vet war a8 Se ce te ee ae ee
DE ee he Ce WN a ee ee ie aw cal ce Tea ais isla ike Uh gh eb al teal a 6
BPUPAR Le oy s a Ce ORT iat i tense a aan Sg giana
FOURTH STAGE LARVAE. Sr ca gious ine eel akeementi ce ike cgilNias Oak giant a ate eee
a ea ces ig cag i gg 11
aureochaeta (Leicester) . .....eee-s Hae set ey Ee
Ce eG ee ee alk ee. le Ee oa ae ee Coe ee
COMMMURTS TLELCESLET sa ke sy es VRE ar are ee ge
eaplex (Theobald) oo. se eek a Gen) elena ae caiat greene 17
COUer 7 aerate aia bE or eee eas ae % gua a ae Week ae 19
demeilloni n. sp. SR gn PAE Ng Cn, te eae) were 21
funerea (Leicester) . Bee Hal coe ce Wane de leh glnas Guia. ober ee Rae Mn 23
indica (Theobald)... .. A ea mee ak egy aR el ee Cs Te 25
macdonaldi Mattingly. .... Dee NE er Uh QCA Sain ar 26.
Re EAA Aa eke ew ew! NR ee ok eee ee 28
POC ee SPO Ma ik ee as Naa ae He MD? ans
PD Se BE ee ee og Se eae ND hig tye ey - 32
Pn ee The ey SO RR a Ree ee ee Se ie ee neh ea he ect le . 33
yer Mattingly... eek. SAP ema mama Ue ON RAMA ema oan ne ta 235
SOU I BO ei ke ei PGR ae eee are agile at a cela ae Gane aa ae ee
BG LAO Ok aig cat eile oes Meek eo weet . 62 eae
PEI ORStES LIOR Oo ia) ace ae eer hea a 6 \pesis OUR Mee
SUBGENUS VATTINGLYIA. Lear eS ew eS age ae ce igh oye: acne tt MD ince
ree eR ICC SEET) i e Seaa es wo Se we ee . 44
a I i Sa GY Who pete ek ea et pe Le eke nigh ae ie ig
PROUHOO Rie BAN aes ek we Bega eee Aa as iad gu Nir ght og cgi 48
PIN Gt te Pe ee ee a a ek a Saat ake gy! Sere ae
LITERATURE CITED. s ose. Be TN get ty See WE ae ee
er ae Wik ge egw a IR Ge: ip hike ce bth wean aie as Unc ae Gh cas aa
DS AR SE ONO er hee ears aan gS ear ana Mer Le
CONTRIBUTIONS TO THE MOSQUITO FAUNA OF SOUTHEAST ASIA, - VI.
THE GENUS HEIZMANNIA LUDLOW IN SOUTHEAST ASIA
By
2
P. F. Mattingly
INTRODUCTION
The Indomalayan species of the present genus were revised by
Mattingly (1957) and the Thailand species by Thurman (1959). Since then a
large amount of new material has come to hand, collected mainly by the staff
of the SEATO Laboratory in Bangkok and the Institute for Medical Research
in Kuala Lumpur. Besides new species this material includes a number of
previously unknown males and early stages. Generic characters have ac-
cordingly required some modification, particularly those affecting the early
Stages which are now very much better known, Lien (1968) has described
three new species from Taiwan. One of these, together with Heizmannia
achaetae and a new species from Thailand, is placed in Mattinglyia Lien
which is downgraded to a subgenus of Heizmannia,
The present paper deals with 19 species from Assam, Burma, Thai-
land, the Andaman Islands, Malaya, Borneo, Indonesia, Vietnam, the
Philippines and Taiwan. Five of these species are new. Others are record-
ed for the first time from particular territories. H. stonei is relegated to
the synonymy of complex and metallica to the synonymy of indica. The Indi-
an species previously described as metallica will now require to be rede-
Scribed under anew name. 4H. funereais removed from the synonymy of
scintillans and the male is described for the first time. One or more of the
new species described here was previously recorded as greenii. The latter
appears to be confined to southern India and Ceylon.
The interesting phenomenon of mouthpart dimorphism (Mattingly,
1957) recorded by Lien (1968) for macdonaldi (as nivirostris Lien), reidi (as
cheni Lien) and faiwanensis is here recorded for communis, reidi and
1 This work was Supported by Research Contract No. DA-49-193-MD- 2672
from the U. S. Army Medical Research and Development Command, Office
of the Surgeon General.
2Department of Entomology, British Museum (Natural History), London SW 7,
England.
2 Contrib. Amer. Ent. Inst., vol. 5, no. 7, 1970
scintillans, It is not known whether the modification of the mouthbrush
hairs to give the fine, filter-feeding or pectinate browsing type is wholly
environmental, as in Opifex (Marks 1958, McGregor 1963), or has become
genetically fixed.
The terminology and general treatment are those adopted by Bram
(1967) with subsequent modifications by SEAMP. The spelling of Thai local~-
ity names is taken from the Official Standard Names Gazetteer #97 prepared
by the Office of Geography, Department of the Interior, Washington, D. C.,
April 1966. Where names are known to represent administrative subdivi-
sions e.g. Changwats, they are italicized. Those locality names which do
not occur in the Gazetteer are spelled according to the labels on the speci-
mens,
GENUS HEIZMANNIA LUDLOW
Heizmannia Ludlow 1905, Canad. Ent., 37:130. Type species: Heizmannia
scintillans Ludlow. Monobasic.
Bolbodeomyia Theobald 1910, Rec. Indian Mus., 4: 31. Type species:
Bolbodeomyia complex Theobald. Monobasic.
Mattinglyia Lien 1968, Trop. Med., Nagasaki, 10: 128. Type species:
Mattinglyia catesi Lien. |
Resembles the New World genus Haemagogus, differing chiefly in
male secondary sexual characters. Features recalling the Sabethini in-
clude the possession by many species of broad iridescent scutal scales and,
in the typical subgenus, a small patch of setulae on the postnotum. The
males have short palpi and, in the typical subgenus, very lightly plumose
antennae, again recalling the Sabethini. The early stages closely resemble
those of Aedes. The larvae of a number of species have a very distinctive
seta C-6, with two very unequal branches, of a kind found elsewhere only in
Aedes subgenus Christophersiomyia. Lien (1968) recently erected a new
genus, Mattinglyia, for achaetae (Leicester) and the closely related cates
Lien from Taiwan. However, thelmae n. sp. from Thailand, shares major
characters of both groups. Mattinglyia is accordingly treated here as a sub-~
genus of Heizmannia.
Generic characters are as follows: |
FEMALE, Small, dark legged species with broad scaled scutum,
frequently showing iridescent colors. Abdominal terga with conspicuous
white or silvery triangular basal lateral markings. Head. Eyes well sepa-
rated; tori with very small, fine hairs and, sometimes, scales on inner sur~
face, appearing bare except under high magnification; clypeus bare except,
sometimes, in complex; vertex with broad, flat, dark scales, small patches
of broad white or silvery scales at the sides and between the eyes and a few
very inconspicuous upright scales low down on occiput; proboscis variable,
long and slender in some species, shorter and stouter in others; palpus
Mattingly: Heizmannia in Southeast Asia 3
short, at most one-fifth of the length of the proboscis; basal joint of antennal
flagellum with small, flat scales; flagellar segments subequal. Thorax.
Scutum with dense covering of broad or moderately broad, flat scales with
dull bronze, matt gold or bright metallic reflections; acrostichal and dorso-
central bristles absent; prescutellars usually absent, at most 1-2; scutellum
with broad, flat scales on all lobes; anterior pronotal lobes unusually large,
almost contiguous in most species; posterior pronotum with broad, flat
scales and with about 2-4 bristles in upper posterior corner; postspiracular
bristles absent in most species; 1-2 lower mesepimeral bristles present;
paratergite with flat, white or silvery scales; similar scales on postspiracu-
lar and subspiracular areas and on sternopleuron and mesepimeron; base of
meron level with, or very slightly above base of hind coxa; postnotum with a
group of small bristles posteriorly, except in Mattinglyia, occasionally mix-
ed with a few broad scales in some species. Legs. Dark except for the fem-
ora; fore and mid femora pale below towards base; hind femur pale on basal
three-fifths or more with dark dorsal stripe usually reaching to base; fore
and mid claws usually simple; hind claws simple in all species. Wing. Dark
scaled; alula with broad, flat, decumbent scales; squama with narrow, hair~
like scales. Abdomen. Tergum I usually all dark; laterotergite white scaled
terga II-VI with conspicuous pale basal lateral markings; some species with
pale basal bands or median spots on some or all of these terga, but with
these marks varying even within the same species; some or all sterna with
dark apical bands. Terminalia, Cerci broad or moderately broad; post~
genital plate with concave or convex margin according to species; [Xth ter-
gum with 2 separate plates (fused in Mattinglyia); VIIth sternum with poste-
rior edge concave, strongly sclerotized and setose; spermatheca triple.
MALE. Resembling the female in general features. Head. Antenna
(Figure 1) subplumose (plumose in Mattinglyia); the 2 terminal joints greatly
elongated except in complex (a single specimen of rezdi with male terminalia
and with antenna of female type is, perhaps, a gynandromorph); palpus short
as in female. Legs. One fore tarsal claw toothed, the other simple in most
species, but both claws toothed in some; mid tarsal claws usually both simple
but one or both toothed in some species. Terminalia. Swollen, conspicuous;
distimere with terminal appendage directed at right angles to the main axis;
inner face of basimere with 1-2 setigerous lobes (claspettes), the more distal
one, as seen in the undisturbed position relative to the base of the basimere,
often with an arm equipped with a falciform or irregularly shaped appendage
(harpago in the original sense of Howard, Dyar & Knab, 1912, p. 69, Fig. 4),
the proximal one often with conspicuous tufts of large, variously modified
setae; inner aspect of sclerotized portion of basimere drawn out into a sub~
apical lobe with 1-2 stout spines (partly modified to form leaflets in
Mattinglyia); aedeagus with toothed lateral plates; apex of basimere with 1-2
rows of modified setae; paraprocts elongated, apically attenuated; [Xth ter-
gum concave with lightly setose lobes.
PUPA. Cephalothorax. Trumpets short, with meatus occupying at
least two-thirds of the length, usually more; the apex expanded in the antero-
posterior plane so that they appear conical in side view, cylindrical when
4 Contrib.Amer. Bate Inst, ’; volo 5): nov: 4. 1970
viewed from in front or behind; seta 1-I well developed, dendritic; 1-II moder-
ately developed with 2-4 branches, sometimes more; seta 5-III-VI normally
single; seta 9-II-VI minute, very inconspicuous, except in aureochaeta and
complex; 9-VII-VUI stout, plumose. Paddle, Oval, usually narrow, with
conspicuous, dark mid-rib and delicate fringe on both borders; seta 1-P long,
Single, sometimes frayed or split.
LARVA. Head. Broader than long; antenna short and straight in
Some species, longer and curved in others, tending to be dimorphic in those
Species with dimorphic mouthbrushes; seta 1-A small, with few or no branches;
mouthbrushes with or without pectinate setae, dimorphic in this respect in
several species; setae 4-C and 7-C multibranched, conspicuous; 5-C with
number of branches very variable; 6-C usually anterior to 4-C, with 2 very
unequal branches in a number of species, of normal type in others; mentum
with 7-11 teeth on either side of central tooth. Thorax and Abdomen. Spines
at bases of pleural setae variously developed; stellate setae absent; integu-
ment of thorax and abdomen without spicules; comb with not more than about
20 scales or spines (except in veidi); siphon without acus, usually about 2.5 -
4,0 times the length of the saddle but sometimes longer (up to nearly 5 times
in some macdonaldi); pecten composed of spines, with or without small basal
denticles, at most about 15 in number, usually much less; seta 1-S long, slen-
der, usually bifid (except in aureochaeta, in which it is trifid or tetrafid), a-
rising at most slightly beyond half way, usually before half way; saddle in-
complete; seta 2-X with 2-4 branches, rarely 5; 3-X single, occasionally bi-
fid; ventral brush with barred area present, usually well developed; seta 4-X
single or bifid, rarely trifid (except in veidi and achaetae);1-2 small precratal
setae usually present; anal papillae large, cylindrical, bluntly rounded or
pointed at tip, showing considerable individual variation.
DISTRIBUTION. Western Himalayas, Southwest India and Ceylon to
Moluccas, Philippines, Taiwan and Korea. Found up to 7000 feet in the
Himalayas.
BIOLOGY. Adults bite man freely, particularly in lowland forest
and, as with Haemagogus, are often more numerous than would be expected
from the number of detectable breeding places. Biting activity is most in-
tense during the hottest part of the day. Breeding places include tree holes
and split or bored bamboos and, in the case of covelli, other containers, such
as split coconuts. Larvae are occasionally found in small ground pools, per-
haps not always in consequence of washing out from other breeding places.
For notes on biology see Macdonald & Traub (1960).
KEYS TO SOUTHEAST ASIAN SPECIES
FEMALES
1, Postnotum bare (Subgenus MATTINGLYIA)...ccccsccccccccceces 16
Postnotum with setulae (Subgenus HEIZMANNIA) ....ccccsacecs 2
2(1). Scutal scales narrow with matt gold reflection..... demeilloni n. Sp.
Scutal scales broad with dull bronze or bright iridescent
Pa eo RS SEG BIOMED Nie COME MSOF RG Na SME ROC RAO 3
3(2).
4(3).
5(4).
6(5).
7(4).
8(3).
9(8).
10(9).
11(10).
12(11).
13(12).
14(13).
15(9).
Mattingly: Heizmannia in Southeast Asia
Scutal scales with dull bronze reflection; posterior pronotum
wholly or mainly pale: Scaled .ccecisccincnss see een eeep.siete os 4
Scutal scales with bright metallic reflection or posterior
pronotum wholly dark scaled or both ...cceccccccecsccceccees 8
Plume scales on vein Ro narrow; postnotum often with
SCaleS. As WEll AS BCUILAS « xs0-0.ka 0. sitien 5:6 6-010 6 min mn meine ea ees 9)
Plume scales on vein Rg broad or very broad; postnotum
PALOLY WIC SCAICS: 6:6 5's diecacecyueieuWiaiele ie Me ieee AieleuarL in aimee wart 7
Hind femur with dark ventral stripe reaching from apex to
TERIA so sh ith ces om: soi he erik tact Sinan te sae te ead a eas taiwanensis
Hind femur entirely pale below unless narrowly dark at tip ...... 6
Proboscis entirely dark; cerci short and square.....ecoeeee cChengi
Proboscis usually with pale spot or streak below at base;
COTCL LONGET ow 4 i:0inie S00 hdielld ae als ek Opies ais ei a ae complex
Anterior pronotal lobes wholly or largely pale scaled... macdonaldi
These lobes pale scaled on anterior border only ....... aureochaeta
Scutal scales with dull bronze reflection; anterior pro-
notal lobes wholly or largely pale scaled .......eeec. communis
Scutal scales with bright, iridescent, metallic reflection;
anterior pronotal lobes various @esooeeseeeaeeseeoeeeweeoeeeeoeneonseeseeoeee¢e& @ ®@ 9
Posterior pronotum entirely dark scaled ....cccscecccccsccces 10
Posterior pronotum wholly or largely pale scaled .........0e0. bs:
Lateral lobes of scutellum with pale scaleS....csescceeee mattinglyi
Lateral lobes of scutetiim Gntirely Gab na ccs wicccicieieenec. cee wt.e 11
Proboscis at least 0.2 as long again as fore femur; cell
Rog at least 2.7 times as long as its stem, often longer..... vreidi
Without this Combimation Of Characierd. 4. viscose ee uc scene + 00 80 12
Hind femur with dark apical band prolonged ventrally,
as well as dorsally, towards base on anterior surface ....... 13
Hind femur pale below, except at tip, and anterior
SUPIACO ss ec ch eka k no 6 ees Ree OAs ARG ACh asa eenaaeaiael MOCETE
Basal lateral pale patches on abdominal terga prolonged
inwards, tapering towards mid line @eeee@aevceaeeoeaoeeaeseeoeeownseoeeeeeee & @ 14
These patches squarer, Truncated. 6... ive cacecceuencedees indica
Anterior pronotal lobes almost entirely dark .....cesccoe Junerea
Anterior pronotal lobes with pale scales extending at
least Some way round anterior border ....ccccceccee scintillans
A spot of broad, whitish scales present at edge of |
scutum above paratergile ..cscoccsccecccecces§ pervsimilis n. Sp,
Scales in this position all dark. wecsscccucseseses | PYOPinaua iso.
Contrib. Amer. Ent. Inst., vol. 5, no. 7, 1970
16(1). Anterior pronotal lobes entirely dark scaled ....... thelmae n. Sp.
17
Anterior pronotal lobes entirely white scaled ...cssccccccccces
17(16). Proboscis about 1.3 times length of fore femur; posterior
pronotum with a few white scales only; wing scales
narrow; lower mesepimeral bristle absent ......ceecc0e- catlesi
Proboscis barely exceeding length of fore femur; posterior
pronotum covered with silvery scales; wing scales broad;
lower mesepimeral bristle present.....ceseccccceeeee achaeltae
The females of proxima n.sp. and scanloni n. sp. are unknown.
—
2(1),.
3(2).
4(3).
0(4).
6(5).
7(6).
8(2).
MALES
Antenna strongly plumose; postnotum without setae; basi-
mere without subapical lobe (Subgenus MATTING L PE OG
Antenna very lightly plumose; postnotum with a tuft of
setae; basimere with well developed subapical lobe
(Subgenus HEI VIANNEA) COp Cees wa wcle ee cee eb Shales cds ee ete 2
Subapical lobe with 2 spines eeoeeveeoeeeaeevaeaeeeeeeoeeaneveee 60608 © @ 3
Subapical lobe with 1 Spine only @eeeeovoeceaeansnoeaoeaoeaeesea~aqsnesne#eoneeeose 0 @ @ 8
Scales of apical tuft, in part, with racket- shaped tips;
appendage of distal claspette bluntly rounded;
POC sels ohn so Ck 6 666 oe Cha Oba eee ves 8 scanloni n. Sp.
Apical tuft and distal claspette OTHET WISE ..cccccvcccccecccvece 4
Last 2 mid tarsomeres with conspicuous scale tufts....... funerea
Mid tarsi without such tufts eeoeoeoesoceeseeoeaeeeaeaeenseaeenesendesoee eee &@ & @
Distimere with greatly enlarged crest; one group
of long setae on proximal claspette fused at base
EGU TOE A COM INTs SNCCL uals ss oie s.c.ed0e meewiciiwcece cee, CHENST
Distimere and ornamentation of proximal claspette
Re Ee aun seselGie, bien wane exe. eusie ehkib eit. o.8 9-001 iniai a wl, Hibie- 6 oe Abele 0:84 6
Appendage of harpago slender, sickle-shaped; apex
of basimere with row of 6 flattened, twisted
Ber ee ea ie hte a ges died ke winiibeienr ahaa iesaie. (elwWanenses
Appendage of harpago broad, crumpled; setae at
apex of basimere only slightly Be ei tas ul casa nails 7
Paraprocts relatively broad ..cceccocccccccccecccsceses COMMUNIS
ParaproctS MuCN NarroweLr .eccccccccccccccccccscves macdonaldi
Distimere deeply cleft, with well developed hair tufts .......0e. 9
Distimere otherwise @eeeaecoececeeeeaeoeaeovneeeeceeeseoaeaeoaeeeeeeee e? © @ 10
9(8).
10(8).
11(10).
12(11).
13(12).
14(13).
15(14).
16(1).
Mattingly: Heizmannia in Southeast Asia 7
Apical tuft with narrow scales only; antennal flagellum
with all segments subequal, as in female......ccseeees complex
Apical tuft, in part, with very broad scales; antennal
flagellum with the 2 terminal segments creatly
CLONGALC Cis: & tis yoiecern-cynieh werk akvn wel etnarerlancls Mekeiethe aelinCeIMeOrnacd
Proximal claspette with expanded, fimbriated
ANPCNGALE: « wie 5: dw avawialld eC isinratede deanna hte ce BRE a he OP eS indica
Proximal claspette without such appendage ....cceccccececccce 11
Proximal claspette with a fan-shaped array of sinuous
filaments almost as long as basimere and distimere
COMDINEGT . 6-0 a-nesiein kebainte:ab-aedaetee erta ee akiten maeeiet Covel)
Proximal claspette othenwise Vic. 2% see enes Banerotatel eee eure lees 12
Distal claspette with a slender, sickle- shaped
DP SNGAGBS pues daeaialley ean eet ata kaaitlates T MRanaM aatictia elie nee galled ale veidi
Distal claspette without such appendage .....ccceccecccocccecs 13
Hind tarsus with a row of erect and suberect scales;
apical tuft with broad scales; distimere complex;
proximal claspette with dense tuft of long, sinuous
setae; distal claspette with 2 tufts of long setae....... scintillans
Hind tarsus without such scales; scales of apical tuft
narrower; distimere relatively simple; long setae
of proximal claspette much less numerous; distal
elaspette. WiEhOUL 1ONG. SEEAS ic is, dis cavekeieta do aie dial eRe OM bie ale 4 AG
Apex of basimere with 2 widely spaced rows of flattened
setae; distimere with convoluted edge......+62. pvopinqua Nn. Sp.
Apex of basimere with a single row of flattened setae
only; distimere with smooth, non-convoluted edge ...-.ceceee 15
Proximal claspette with several dark, sinuous setae
and with all setae of only moderate length ........ pvoximan.Sp.
Proximal claspette without conspicuous, dark setae,
with a tuft of very long, sinuous Setae ..ceseoee Persimilis n. Sp.
Posterior pronotum covered with silvery scales;
apex of basimere with 2 apically expanded leaflets
and a patch of flattened, fimbriated setae.....c.cececceee achaetae
Posterior pronotum with a few white scales only;
apex of basimere with 2 relatively narrow leaf-
lets and. .a patch:ebhaireqew.d5s Hedy sake aswsdeenrcineee Clbest
The males of demeillonin. sp., mattinglyi and thelmae n.sp. are unknown.
aay
e
2(1).
3(1).
4(3).
(4).
6(5).
7(5).
8(7).
9(8).
10(9).
Contrib. Amer. Ent. Inst., vol. 5, no. 7, 1970
PUPAE
Seta 5 on segment V of abdomen much less than half
as long as segment Vi @eeoece@aeaeeoeoeeospnecese@escenvneesonoesgeeesesesvseeee#e#@g 2
This seta at least three-fifths as long as segment VI .....ccece000 3
seta 5 on segment IV very short; seta 9 on segments
II-VI a short but conspicuous dark spine ......ee2e. aurveochaeta
Seta 5 on segment IV much longer; seta 9 on II-VI
minute, almost invisible under low power .....ccceeeee» a&Chaetae
Seta 5 on segment VI normally at least 1.2 times the
length of the following segment; paddle fringe very
long and coarse @eccoco@eeeseseaeeseeeseoaeoeseeseeseeaneaeeeeneaegvnee @ 80286 8 @ @ reidi
WitDouE this Combination Of Characters ¢ . ss sos ss «0 6:6 0.0 0 00:00 4
seta 9 on segments II-VI a conspicuous dark spine;
seta 5 on 0 long, stout and dark, resembling 2
AG OH GOiMie SEPRICNE case's selec ve esscaeesckecseccvcces. COMPLEX
Without this conibination of characters vicsecceseccsrcceccecee 9)
Seta 1 on segment II with abundant secondary branching;
seta 6 on cephalothorax less than half as long as
seta eoeeovaees@eaeeeoeoeaocpeeae@ot CGeseegeeeaee@sopeoeaseoaeeoweeoescseseeeoeeeeoe se 6
Without this’ combination of Characters Viiissece ceeds acace ceewe %
Paddle generally narrower; 1.6 - 2.0 times as long
as broad . eeoeog@eseoeeeeeeseeeé ee ee CO Oe eeoo@eceodoeeeoee ae @ @ se macdonaldi
Paddle broader: 1b 9 times as long as broad. BY communis
Seta 5 on segment VI at most two-thirds as long as
segment VII .... @eeoee?s 8 e @®eosceeseeeeseoecscse?@ 8
This seta normally at least three- fourths as ‘long as
segment VII . @eoeoeooeocesv ogoe08e268@# 0602082 G8GeCGePeKGKGeeKBHeKQKSee#eBBKHBeEKGCAKABeBE 11
seta 9 on segment II-VI relatively long, dark,
CORSOICUOUS jap cioceio ew eia dias Pana Min ews ed Mote ted Cen LT
This seta minute, colourless, MET WICONBDICUOUS wick a cewek wees
Seta D5 on segment IV almost as long as or longer
than segment V . °eecg e828 e686 8 Ge eB e @e9@eesoeeecseoeoeseeeoeseeeeeesgeeeoesd ge @ eoee38 @ 10
This seta much shorter than segment We he Se OG 9 thelmae Nn. Sp.
Paddle long and narrow, at least twice as long as
Ce ea ee Ie Sik ede eke baw dew e das eoea er PYOXIMA TE Spi
Paddle shorter, at most about 1.8 times as long
SES Oe ee OPO CUID bebe o ee kA ee Aes RRO be ee eelenae ei 12
11(7).
12(10).
13(12).
Mattingly: Heizmannia in Southeast Asia 9
Male genital sac with narrow apical horns ....... pevsimilis n. Sp.
? propinqua nN. Sp.
Male genital sac usually with apices bluntly
PoUndeS by TTS ee ha eke ct kaleen be Ko ees Gr rire
Paddle fringe unusually short and scanty; known
only from Burma and Thailand .....ccescceccce demeilloni n. Sp.
Paddle fringe not unusually short and scanty;
known only from Taiwan ..ceccoccccsccccveseccesescccsescees uo
Paddle oval, pointed apically ..ccccscccsesccccceseece laiwanensis
Paddle pear-shaped with apex flattened . tes Se ee Noe ee ne | RE Oe
The pupae of covelli, indica, mattinglyi, funerea and scanloni n. Sp. are
unknown. |
2(1).
3(2).
4(3).
5(3).
6(5).
FOURTH STAGE LARVAE
Setae of ventral brush with numerous branches;
comb of at least 30 scales with uniformly
fringed AVEOX o'ssic.4 + iain gued MUN ee 4 Aa ARR eek Oe ee
Without this combination of characters .....ccccces bee eK SR 2
All or most setae of ventral brush with at least 3
branches; pecten teeth minute...cccccccecscecccceccese achaetae
catest
Without this. combination ol characters ani cash oes oe Wp a eee ee es
All comb teeth with uniform apical fringe ....cocccccceccecvcccs 4.
Comb teeth, at least in part, with hypertrophied
median denticle eceoeaoceaeeceswsaevsveoeavn00e0@e2000@02@02080280008000090809080 0 @ 5
Antenna smooth; seta 1-A single or bifid; pecten
teeth stouter, the distal ones, at least, without
secondary Clentle ei y.nete:sitieid-s on awnd Ded see name ee namie) COMenTe
Antenna spiculate; seta 1-A with 2-4 branches; distal
pecten teeth with conspicuous secondary demticlem caus: euieaits ae
Comb consisting entirely of spines with hypertrophied
WACOIST. COREG IG ike pe io ede een Od Cae Re Ras ee 6
Comb of mixed spines and uniformly fringed scaleS ...ecccocees 9
Comb spines with median denticle very short; antenna
shore. and StPaigt 066 de wehe ee bas case hae aek eee eee aureochaeta
Without this combination of characters ....ccccceccccccecccece 7
10
7(6).
8(7).
9(5).
10(9).
11(9).
12(11).
13(4).
14(13).
Contrib. Amer. Ent. Inst., vol. 5, no. 7, 1970
Spines of bases of pleural setae very Small .....cccccceccccece 8
These Spines laree, CONSDICUCUS .4icsescecsedcesdes PYOXIMG Ns SPs
Antenna long, curved; pecten well developed...... demeilloni n. sp.
Antenna short, straight; pecten teeth few, small..... thelmaen.sp.
Head seta 6-C usually with 2 or more approximately
SOOO MCS: 10. is tuet ateue fase as sks ekebarenel thcreaibavantie towrk, ace: le wledelmun a « 10
This seta single or with 2 very unequal branches ....ccccoeccee Ll
Head seta 6-C usually with 3 or more branches;
Siphon at least 4.5 times as long as its breadth
Fee te len eta eth enue e pain ace lee in we Biels Oa ee Sead eee MUACKONAIAL
This seta normally bifid, rarely trifid; siphon
Cle aor rer WG ES. fags on eke tke oo aa oh 6% ose 88 communis
All comb teeth with median denticle at most very
feebly developed; most pecten teeth with con-
Spicuous secondary denticles; mouthbrushes with
pectinate setae (? alwayS) ..cscccccvccsecescess Propingua Nn. Sp.
Without this combination of characterS ...cccccccccccvcccecees 12
Pecten teeth usually smooth or almost so, at most
1-2 of the more distal teeth with conspicuous
CN Ts wee ane a vase oid ba Ak Kia ol yb S ecouesk © hia as ane.e 4.8 1, HO UECL LS
funerea
At least 3 teeth with conspicuous denticles ......ccee0ee persimilis
Comb of 7-14 fringed scales of more or less
uniform size ee@ee@ocoaooaoeesesoeaeeaeeaeosaaoaeeeogeeoeoevsaeasedveees @ @ é @ @ 7 covelli
Comb scales more numerous @eeaeoeo@*eaoeaescoaoeeeeeeoeasoeaenvneeeeeeesseé eee 14
Comb with 2 groups of scales, the dorsal scales
much longer than the ventral; distal edge of
SHGALE WILD CONGDICUOUS SDINGS .. ces bc vie sets epee seececcs, CHERR?
Comb scales more or less equal in size; distal
edge of saddle with fine spicules only ......eccccecee ldiwanensis
The larvae of indica, mattinglyi and scanlonin. sp. are unknown.
Mattingly: Heizmannia in Southeast Asia 11
SUBGENUS HEIZMANNIA LUDLOW
Heizmannia Ludlow 1905, Canad. Ent., 37: 130. Type species: Heizmannia
scintillans Ludlow. Monobasic.
Bolbodeomyia Theobald 1910, Rec. Indian Mus., 4: 31. Type species:
Bolbodeomyia complex Theobald. Monobasic.
Characterized by the presence of postnotal setae, simple fore and
mid tarsal claws of females of most species, subplumose male antennae and
well developed subapical lobe of male basimere bearing 1 or 2 stout spines.
HEIZMANNIA (HEIZMANNIA) AUREOCHAETA (LEICESTER)
(Figs. 2, 9;3, 02 terminalia, * claws; 4, pupa, larva)
Dendromyia aureochaeta Leicester 1908, Cul. Malaya, 255 (¢).
Resembles complex, macdonaldi and communis, and differs from
other species, in the subdued, bronzy reflections from the scutal scaling.
Differs from these 3 species in the characters given in the key. The male ter-
minalia recall those of complex, from which, however, they differ in many
details. The stout, unbranched seta 6-C of the larva also recalls complex but
the comb and pecten teeth are very different in the 2 species. The pupa re-
sembles that of complex in the well developed seta 9-III-VI and that of achaelae
in the reduced seta 5-V. This is, however, a distinctive species in all stages,
unlikely to be confused with any other. Principle characters are as follows.
FEMALE. Head. Top of head with broad, flat, dark scales and with
flat, creamy scales at sides below, the latter not visible from above; a narrow
border of small, flat, white scales round the eye margins, sometimes reach-
ing to mid line; a triangular or diamond-shaped patch of snow-white scales be-
tween the eyes which are widely separated; basal joint of antennal flagellum
with small, white scales on inner edge; proboscis short and relatively stout,
about nine-tenths of the length of the front femur or slightly less; clypeus bare;
palpus about one- seventh to one-fifth of the length of the proboscis. Thorax.
Scutal scales with dull greenish bronze reflections; a few pale scales some-
times present immediately in front of mid lobe of scutellum and at apex of
scutum; prescutellar bristles absent; scutellum dark except for a few broad,
pale scales at apex of mid lobe in unrubbed specimens; upper surface of ante-
rior pronotum dark except along the anterior border; posterior pronotum with
broad, pale scales and with some broad, dark ones in upper posterior corner;
12 Contrib. Amer. Ent. Inst., vol. 5, no. 7, 1970
pleuron as described for achaetae except that the scales mostly have a creamy
tinge; postnotum with a conspicuous patch of pale, very narrow scales or scale-
like bristles and occasionally 1 or 2 broad scales. Wing. Plume scales on
veins Ro and Rg very broad; cell Rg about 3.5 - 4.5 times as long as its stem.
Legs. ftind femur pale anteriorly and posteriorly on about the basal three-
quarters to four-fifths with a broad, dark dorsal stripe reaching to base;first
hind tarsomere pale below; all tarsal claws simple. Abdomen. Tergum I
dark; remaining terga dark above with basal lateral creamy white patches
prolonged backwards, ventrally, nearly to the posterior border on II, HI and
VI and sometimes also on IV and V, occasionally with median basal pale spot
on IV-VII. Sterna II and III wholly or largely pale, IV-VI broadly dark at
apex, VII dark. Terminalia. Postgenital plate small, convex, evenly round-
ed; postatrial plates moderately to strongly sclerotized.
MALE. Head, Antenna with the 2 terminal segments elongated; rest
of flagellum much as in female. Legs. Fore claws each with a large tooth
near base; one mid claw with a large, the other with or without a small tooth
near base. Terminalia. Distimere deeply cleft, with tufts of hair resem-
bling that of complex but with a short arm arising from outer edge near base;
apex of basimere with a row of flattened, striated leaf-like setae and 1-2
similar setae between this row and the apex; subapical lobe with a long, sinu-
ous spine; claspette greatly reduced, the distal claspette @cl) with short hairs
only, without harpago, the proximal claspette (cl) with a few long, sinuous
hairs; paraprocts each with 2 short, thumb-like processes at apex; Xth ster-
num membranous, pilose at base; aedeagus without special features; IXth
tergum with 2 shallow lobes, each with about 4 small bristles; [Xth sternum
sclerotized basally and around edges and with a medium sclerotized area
bearing several short bristles.
PUPA. Cephalothovax, Seta 6-C shorter than 7-C, single; 7-C
single to trifid. Abdomen. Seta 1-II small with 2-4 simple or lightly plu-
mose branches; 5-II slender, very delicate, single; 9-II-VI small and slender,
but less so than in most other species, easily visible under low power; 5-IV-
VI very small, less than half the length of the following segment, single, oc-
casionally split distally; 9- VII and 9-VIII both short, with 1-3 and 1-4 branches
respectively. Paddle. Variable, often broader than usual for the genus,
about 1.2 - 1.7 times as long as greatest breadth, with fringe formed by a
double row of fine, hair-like spicules; seta 1-P usually single, sometimes bi-
fid; male genital sac triangular, the apical horns only slightly differentiated
from the remainder.
LARVA. Head, Antenna short, cylindrical, smooth or almost so;
seta 1-A single or bifid, often lightly plumose; inner mouthbrush setae pecti-
nate in all cases; seta 5-C with 3-8 branches, usually 5-7; 6-C single or bifid,
when single often swollen just beyond base. Abdomen. Comb of 7-13 spines
(cs) with short, stout median denticles and conspicuous basal fringe, in 1-2
rows; siphon about 2,3 - 3.0 times the length of the saddle (about 2.5 times
its breadth at base); pecten of 3-8 small but stout spines (pi) which usually
have 1 or more basal denticles but are occasionally simple; seta 1-S with 3-5
branches, equal to or shorter than diameter of siphon at point of attachment,
Mattingly: Heizmannia in Southeast Asia 13
usually arising beyond the terminal pecten tooth but occasionally between this
and the subterminal tooth; saddle with distal edge very lightly spiculate; seta
1-X long, stout, plumose, bifid; 2-X bifid; 3-X single; anal papillae bluntly
rounded, between 1.5 and 2.0 times the length of the saddle; ventral brush
with 8-10 setae, the 6 most distal bifid, the rest with 3-5 branches.
TYPE DATA. Lectotype female, on pin, in British Museum; type
locality: Jungle 6 miles from Kuala Lumpur, WEST MALAYSIA.
DISTRIBUTION. Specimens examined: THAILAND, Nakhon Si
Thammarat, Banna Nabon 4 females, Khao Nawua Banna 1 female, Khao
Luang Banna 1 female; Phrae, Rong Kwang 1 female; Loei, Phukaoung 1 fe-
male, Pat Meeung Mountains 1 female; Ranong, Khao Chatri 2 whole larvae;
Tak, Khao Salak Phra 1 female. CAMBODIA, Pichnil 2 females. WEST
MALAYSIA, Selangor, Kuala Lumpur, Ulu Gombak, Ulu Langat and Pahang
road 30 females, 11 males, 17 larval and pupal skins, unnamed localities 2
females. Other Distribution. Recorded by Causey (1937) from Bangkok, Bon
Pin and Chiang Muang, THAILAND.
TAXONOMIC DISCUSSION. This is a very distinct species, unlikely
to be confused with any other. It is, perhaps, most closely related to complex,
which it resembles in the male terminalia. One specimen from Cambodia has
very small median basal pale spots on terga III-VI.
BIOLOGY. Said to bite man in daytime, in dense shade only (Causey
1937). Adults have been taken on man in forest, bamboo forest and a rubber
plantation. Larvae have been found in bored bamboos in Malaya (Macdonald
& Traub 1960) and in a flood pool in a bamboo grove at 2900 ft. in Thailand.
HEIZMANNIA (HEIZMANNIA) CHENGI LIEN
(Figs. 5,0°92;6,09 terminalia, ¢ claws;7, pupa, larva)
Heizmannia chengi Lien 1968, Trop. Med., Nagasaki, 10: 137 (o'*, 9*, P*, L*),.
This species closely resembles complex in external features of the
female. The male antenna and terminalia are entirely different, however, the —
latter showing an interesting resemblance to those of covelli in the ornamen-
tation of the proximal claspette.
FEMALE. Head. Top of head with narrow pale line bordering eyes,
triangular pale patch between eyes; eyes comparatively close together behind;
basal joint of antennal flagellum with conspicuous pale scales on inner side;
palpus and proboscis dark; proboscis about 1.0 - 1.1 times the length of fore
femur; palpus about one-seventh the length of proboscis. Thovax. Scutal
scales with dull bronze reflection; a narrow line of pale scales round anterior
border; pale scales on paratergite and a few in front of wing root; a small spot
of pale scales on mid lobe of scutellum; anterior pronotal lobes with a pale
spot in anterolateral corner and a few inconspicuous pale scales on anterior
margin; posterior pronotum with patch of well developed bristles and some
broad, pale scales. Wing. Plume scales narrow; anterior fork cell about
2.5 times as long as its stem. Legs, Anterior surface of hind femur pale
14 Contrib. Amer. Ent. Inst., vol. 5, no. 7, 1970
below on about the basal nine~tenths with dark dorsal stripe reaching base;
fore and mid tarsal claws simple. Abdomen. Laterotergite pale scaled;
terga II and III with basal lateral pale patches prolonged ventrally to posterior
border of segment; terga IV-VI with similar patches but these not reaching the
posterior border of the segment. Terminalia. Cerci shorter and squarer and
postgenital plate broader than in complex.
MALE. Head. Antenna with the two distal flagellomeres greatly e-
longated; verticillary hairs on all flagellomeres short and scanty. Legs. One
fore tarsal claw simple, the other with a large, straight tooth near base; mid
claws simple or one with a straight tooth arising near base (see under Taxo-
nomic Discussion). Tervminalia, Distimere with a rounded, transparent
crest and a short arm arising from the base of this; apex of basimere with a
row of 4 only slightly flattened setae; subapical lobe with 1 stout and 1 more
slender, twisted spine; distal claspette with well formed harpago (#) with fal-
ciform appendage; proximal claspette (cl) with a bunch of long, sinuous, flat-
tened setae fused at base to form a transparent, striated sheet; paraprocts
each with a thumblike process and 2 small teeth at tip; [Xth tergum with well
separated lobes each with 4-6 fine hairs; [Xth sternum lightly but extensively
sclerotized with numerous scales and 1-2 small setulae near tip.
PUPA. Cephalothorax. Seta 6-C shorter than 7-C, single; 7-C
single or bifid. Abdomen. Seta 1-II slender with 2-4 branches; 5-II short,
single, somewhat stouter and darker than usual; 9-II-VI short but dark and
relatively conspicuous; 5-III-V single; 5-IV and 5-V slightly shorter than the
following segment; 5-VI very short, single or bifid; 9- VII single; 9- VIII single
to trifid; male genital sac with apical horns well differentiated. Paddle.
Fringe relatively short and delicate; seta 1-P single.
LARVA. Head. Antenna relatively long, slender, curved, lightly
spiculate; seta 1-A bifid; inner mouthbrush setae delicate, nonpectinate; seta
5-C with 13-14 branches; 6-C bifid or trifid with 1 branch usually longer and
stouter than the other(s). Abdomen. Comb with about 22-27 uniformly
fringed scales (cs), the dorsal group much larger than the ventral; siphon
about 4 times as long as the saddle (about 3.5 times its breadth at base); pec-
ten with 6-11 teeth (pt), the proximal ones longer, simple, the distal ones
shorter, simple or with a secondary denticle at base; seta 1-S long, single;
saddle with distal edge strongly spiculate; seta 1-X single; 2-X bifid; 3-X
single; anal papillae very long, bluntly rounded at tip; ventral brush with 6
long setae in barred area and 2 short precratal ones; 1-2 of the long setae and
sometimes 1 or 2 of the short ones bifid, the others single.
TYPE DATA. Holotype male and allotype female, with associated
larval and pupal skins, in Taiwan Provincial Malarial Research Institute,
Taipei; type locality: Peiyuan, Tungho Township, Taztung, TAIWAN,
DISTRIBUTION. Specimens examined: TAIWAN 1 male, 2 female
paratypes, with associated larval and pupal skins, from the type locality.
THAILAND, Chumphon, Muang 1 male; Khon Kaen, Padong Larn 2 females,
1 set of male terminalia; Nakhon Ratchasima, Ban Wang Ta Khian 1 male with
associated pupal skin.
Mattingly: Hezzmannia in Southeast Asia 16
TAXONOMIC DISCUSSION. Taiwan and Thailand specimens agree
well except that the only Thailand male examined has both mid tarsal claws
simple whereas the male from Taiwan has one of them toothed. Too little is
known regarding the extent of variation in this character to assess its signifi-
cance. On present evidence I would not feel justified in describing the Thai-
land form as a distinct species or even subspecies.
BIOLOGY. The type series was bred out from a tree stump, with
Orthopodomyia anopheloides and Tripteroides bambusa, on a forested hill at
an altitude of 260 m. The male from Nakhon Ratchasima was bred out from a
tree hole. The male from Chumphon was caught in a light trap.
HEIZMANNIA (HEIZMANNIA) COMMUNIS (LEICESTER)
(Figs. 8, 2; 9,0? terminalia, claws; 10, pupa, larva)
Dendromyia communis Leicester 1908, Cul. Malaya, 254 (°, 9).
This species and macdonaldi form a closely related species pair.
There seem, however, to be constant differences in all stages. Relationships
to other species are obscure.
FEMALE. Head, Top of head with broad, flat, dark scales and flat
white scales at sides, not visible from above; no pale scales round eye mar-
gins; a large diamond-shaped patch of silvery white scales between eyes; eyes
well separated; basal joint of antennal flagellum with small, wholly or partly
dark scales; proboscis short and stout, about nine-tenths of the length of the
fore femur; clypeus bare; palpus about one-fifth of the length of the proboscis.
Thorax. Scutal scales with dull, bluish bronze reflection; prescutellar bris-
tles absent; scutellar scales all dark; upper surface of anterior pronotal lobes
covered with broad, white scales, sometimes with a few dark scales posteri-
orly; posterior pronotum with dark scales only; paratergite with dark scales
and some white ones along lower edge; posterior edge of sternopleuron with a
stout bristle and, usually, 1-2 fine hairs in addition; pleuron otherwise much
as in achaetae; postnotum with a patch of bristles, mostly rather pale. Wing.
Plume scales on veins Rog and Rg broad; cell Ro about 3.5 times as long asits
stem. Legs. Hind femur pale on about the basal three-fifths and, sometimes
obscurely so for some distance beyond this, with dark dorsal stripe reaching
to base; all tarsal claws simple. Abdomen. 'Tergum I dark; terga II-VI dark
above with basal lateral white patches prolonged backwards ventrally only on
II and, sometimes, slightly on III; tergum VII all dark or with narrow, in-
complete, basal white band; sterna with dark apical, and pale basal bands,
the latter often more yellowish in color than the basal lateral tergal patches.
Terminalia. Postgenital plate bilobed; otherwise much as in aureochaeta.
MALE. Head, Antenna much as in the 2 previous species; probos-
cis distinctly longer and more slender than in the female. Legs. One fore
tarsal claw with a long, slender tooth arising from base; the other simple.
Mid tarsal claws both simple. Terminalia. Distimere relatively simple,
with large terminal appendage; apex of basimere with a double row of setae
16 Contrib. Amer. Ent. Inst., vol. 5, no. 7, 1970
which are slightly, but not very conspicuously, flattened; subapical lobe with
2 unequal spines; distal claspette with a harpago (h) with broad, densely pi-
lose stem and expanded transparent, membranous appendage; proximal clas-
pette (cl) with a bunch of long, twisted setae and 2 groups of shorter, flat-
tened setae; paraprocts very lightly sclerotized, elongated, broadened to-
wards apex; IXth tergum with lobes lightly sclerotized, well separated, each
with 3-7 fine hairs; [Xth sternum smaller than usual, strongly sclerotized,
with a row of about 4 small bristles at apex.
PUPA. Cephalothorax. Seta 6-C single (bifid on one side in one
specimen), shorter than 7-C which has 1-4 branches. Abdomen. Seta 1-II
conspicuous, Semi-palmate with short stem and considerable secondary and
tertiary branching; 5-II small, very slender, single; 9-II-VI small, delicate;
5-IV and 5-V from somewhat shorter than to somewhat longer than the follow-
ing segment, single or bifid; 5-VI single, about one-third to three-fifths of
the length of the following segment, occasionally split towards tip; 9- VI and
9-VIII single to trifid and trifid to pentafid respectively. Paddle. From about
1.5 to 1.9 times as long as its greatest breadth with double fringe of fine
hairs; seta 1-P usually bifid, rarely single or trifid on one side; male genital
sac with apical horns usually, but not always, well differentiated.
LARVA. Head, Antenna dimorphic, shorter in those larvae with
pectinate mouthbrush setae, longer in those without, lightly spiculate both
basad and distad of seta 1-A; this seta bifid to tetrafid, simple or lightly plu-
mose; mouthbrushes dimorphic, either with simple hairs only or with the in-
ner setae strongly pectinate; seta 5-C with 3-10 branches; 6-C usually bifid,
occasionally trifid on one side. Abdomen. Comb a mixture of 14-28 scales
(cs), with uniform apical fringe, and spines each with a hypertrophied median
denticle; siphon about 3.1 - 5.0 times the length of the saddle (about 3.0 - 7.0
times its breadth at base); pecten of 4-14 spines (pf), all or most of them usu-
ally with well developed secondary denticles and expanded at base; seta 1-S
with 2-4 branches, approximately equal in length to the breadth of the siphon
at point of attachment, variable in position, often arising at a considerable
distance beyond the terminal pecten tooth; saddle with apex spiculate; seta
1-X long, lightly plumose, single or bifid; 2-X with 2-4 branches; 3-X single;
anal papillae broad, bluntly rounded at tip, the ventral pair somewhat smaller
than the dorsal; ventral brush with about 8 setae which are mostly bifid (most-
ly trifid in one larva), 1 or 2 sometimes single, the 2 smaller proximal setae
without barred sockets.
TYPE DATA. Hololectotype male and allotype female, on pins, in
British Museum; type locality: The Gap, Selangor, WEST MALAYSIA.
DISTRIBUTION. Specimens examined: THAILAND, Chanthaburi,
Chaung Khao 17 females, 2 males, 13 larval and pupal skins, 3 pupal skins,
22 whole larvae; Phangnga, Nam Tail female, 1 male, 1 pupal skin, Tang
Mai 2 males, 2 pupal skins; Ranong, Ban Chatri 8 females, 1 male, 5 larval
skins, 9 pupal skins, 3 whole larvae. WEST MALAYSIA, Selangor, The Gap
3 females, 2 males; Fraser's Hill 1 female, Ulu Langat 3 females, Ulu
Gombak 16 females,11 males, 18 larval and pupal skins; Pahang, Raub 1 fe-
male; Perak, Cameron's Highlands, 2500 feet 1 female. Unnamed Malayan
Mattingly: Heizmannia in Southeast Asia Be
localities 6 females. Other Distribution. Recorded by Brug & Bonne-Wepster
(1947) from INDONESIA, Sumatra and by Borel (1930) from COCHIN CHINA
(but see under covelli).
TAXONOMIC DISCUSSION. Adults of both sexes can be separated
from those of macdonaldi by the ornamentation of the posterior pronotum.
Individual larvae and pupae may present difficulties but almost all those seen
by me have been readily separable on the characters given in the keys. The
larval mouthbrush dimorphism has been discussed above (p.16). Nearly all
the Malaysian larvae of the present species seen by me have browsing mouth-
brushes but in all except one of the Thailand larvae they are of the filter feed-
ing type.
BIOLOGY. Recorded as attacking man in forest. Breeding places
mainly bored bamboos with small or moderate sized holes (Macdonald &
Traub 1960).
HEIZMANNIA (HEIZMANNIA) COMPLEX (THEOBALD)
(Figs. 11, 9; 12, *¢ terminalia, o claws; 13, pupa, larva)
Bolbodeomyia complex Theobald 1910, Rec. Indian Mus. 4: 31 (¢, 9).
Heizmannia stonei Mattingly 1957, Cul. Mosq. Indomalayan Area, 2: 51 (9*).
NEW SYNONYMY.
Closely resembles chengi in external features but separable, in most
cases at least, by the presence of a pale ventral spot or streak at the base of
the proboscis. The female terminalia are also distinctive and the male termi-
nalia are entirely different. Thailand specimens often have scales on the
clypeus as well as on the postnotum. H. complex from Burma and Assam ap-
parently lack these from both clypeus and postnotum. 4H. stoneéi was provi-
sionally recognized as distinct by reason of the presence of these scales.
However, the male terminalia proved to be identical with those of complex.
I have therefore relegated it to the synonymy.
FEMALE. Head. Eyes with a more or less complete border of
broad pale scales, closer together than in most other species; basal joint of
antennal flagellum pale scaled; proboscis variable, from longer than to some-
what shorter than fore femur, with a pale spot or streak below at base; palpus
about one-sixth the length of the proboscis or slightly less; clypeus bare or,
in some Thailand specimens, with broad, flat, white scales. Thorax. Scu-
tum covered with dark bronzy scales with subdued metallic reflection; broad
pale scales usually present around anterior border, in front of wing roots and
in front of mid lobe of scutellum; prescutellar bristles absent; mid lobe of
scutellum with apical pale scales; pale scales also frequently present on lat-
eral lobes; upper surface of anterior pronotal lobes pale scaled around whole
of anterior border, dark behind; posterior pronotum entirely pale scaled;
postspiracular area with 1-2 pale bristles; pleuron otherwise as described
for the previous species; postnotum with a small patch of bristles and, fre-
quently, some broad, pale scales in addition. Wing. Plume scales narrow
18 Contrib. Amer. Ent. Inst., vol. 5, no. 7, 1970
or very narrow; cell R»y about 2.3 - 2.8 times the length of its stem. Legs.
Hind femur pale on about the basal four-fifths to nine-tenths of the anterior
surface with dark dorsal stripe reaching to base; all tarsal claws simple.
Abdomen. TergalI and II all dark or with median basal white spot; terga II to
VII all dark above or one or more of these with a wholly or partly complete
white basal band; all terga with basal lateral white patches which tend to be
prolonged posteriorly at the outer edges; sternum II entirely pale; sternum
III pale or with dark apical band; remaining sterna with dark apical bands.
Terminalia. Much as in aureochaeta;IXth tergum without setae (? always).
MALE. Head, Antenna with all joints subequal, as in female;
verticillary hairs short and scanty. Legs. One fore tarsal claw simple, the
other with a large tooth at base; mid tarsal claws both simple. Terminalia,
Distimere deeply cleft with complex ornamentation, much as in aureochaeta
but without the small arm near base; apex of basimere with a row of 5 flat-
tened, striated setae; subapical lobe with a single slender, twisted spine;
distal claspette @cl) with a slender, very transparent arm, expanded at tip;
proximal claspette represented by a small, well sclerotized lobe without
bristles; paraprocts lightly sclerotized with thumb-like projection at apex;
IXth tergum with lobes closely approximated, each with about 6-9 fine hairs;
IXth sternum sclerotized at sides and with sclerotized median arm bearing a
number of bristles.
PUPA. Cephalothorax, Seta 6-C long, stout and single (bifid on one
side in one specimen), much longer and stouter than 7-C which is single or
trifid. Abdomen. Seta 1-II with 2-5 branches (7 and 8 in the only Thailand
specimen); 5-II long, stout, single (bifid on one side in one specimen), very
conspicuous; 9-II-VI usually long and conspicuous; 9-VII single (sometimes
split distally); 9-VIII single to trifid. Paddle, About 1.8 - 2.0 times its
greatest breadth, with double fringe of fine hairs; seta 1-P slender, single;
male genital sac with apical horns not very well differentiated.
LARVA. Head. Antenna short, straight, smooth; seta 1-A single or
double; mouthbrushes with inner setae pectinate in all cases; seta 5-C single,
usually expanded just beyond base, often markedly so; 6-C with 1-3 branches
(4-8 in Thailand larvae), Abdomen. Comb of 8-18 dark, finely fringed scales
(cs), the majority, usually, rather long and narrow; siphon about 4.0 - 5.0
times the length of the saddle (about 3.0 - 4.0 times its breadth at base; pecten
of 1-5 (usually 3-4) spines #7), all or most of them, usually, with secondary
denticles minute or absent; seta 1-S bifid, arising well beyond the distal pecten
tooth; saddle with apex strongly spiculate; seta 1-X long, lightly plumose, bi-
fid or, rarely, trifid; 2-X with 2-3 branches, rarely 4; 3-X single; anal papil-
lae broad, cylindrical, bluntly rounded at tip, the ventral pair slightly shorter
than the dorsal; ventral brush with 8 setae, the 2 most anterior with 2-5
branches, the other with 2-3.
TYPE DATA. H. complex (Theobald), hololectotype male terminalia
and wing, on slides, in the British Museum; remainder of this specimen, on
pin, in Indian Museum; allotype female, on pin, in Indian Museum; type local-
ity: Dawna Hills, LOWER BURMA. JH. stonei Mattingly, type female, onpin,
in British Museum; type locality: Ulu Langat, Selangor, WEST MALAYSIA.
Mattingly: Heizmannia in Southeast Asia 19
DISTRIBUTION. Specimens examined: INDIA, Assam, Nongpoh 2
females; Haflong 1 female. BURMA, Dawna Hills 1 female, 1 male. THAI-
LAND, Chiang Mai, Doi Suthep 1 female; Nakhon Nayok, Ka- Ang Waterfall
1 female, 2 whole larvae, 1 larval and 1 pupal skin, Sariga Village 1 female;
Nakhon Si Thammarat, Khao Luang Banna 1 female; Udon Thani, Muang 5 fe-
males, Ban Kut Ling Hyou 1 female; Chon Buri, unnamed locality 2 females;
Phrae, Rong Kwang 2 females; Lamphun, Kang Arb Nang 2 females; Ubon,
Chong Mek 1 female. WEST MALAYSIA, Selongor, Ulu Gombak 19 females,
5 males, 15 larval and pupal skins, Ulu Langat 1 female, The Gap 3 females.
VIETNAM, Cheo Reo 1 female, Cam Ranh 11 females. CAMBODIA, Kirirom
1 female, Pichnil 1 female. Other Distribution. Recorded from locality be-
tween Bon Pin and Chieng Molang (? = Bon Chiang Muang), THAILAND by
oa? (1937). Recorded by Borel (1930) from Terres Rouges, COCHIN
CHINA.
TAXONOMIC DISCUSSION. Specimens from Tad Muay Falls, Phrae
Province, Thailand, provisionally recorded by Thurman (1959) as stonei are,
in fact, achaetae. Her records of complex are here confirmed. Borel's de-
scription and figures of complex from Indochina differ in some small particu-
lars from the above but it is significant that he described the male antenna as
having all segments equal, a very unusual feature otherwise known to me only
in the present species and in an aberrant male of reidi (p.35). His drawing of
the male terminalia, though rough, also agrees quite well with the present
species as does his description of the rather characteristic larval comb
scales. I am inclined, therefore, to accept his record, particularly as there
are females agreeing well with this species, from Vietnam, in the U. S.
National Museum.
BIOLOGY. Recorded as attacking man in forest. Causey (1937) re-
cords it as collected on a horse. Larvae have been found in tree holes in
Malaya (Macdonald & Traub 1960) and in bamboos in Thailand.
HEIZMANNIA (HEIZ MANNIA) COVELLI BARRAUD
(Figs. 14, °9;15, ¢? terminalia, « claws; 16, larva)
Heizmannia covelli Barraud 1929, Indian J. med. Res. 17: 265 (c'*, 9).
This species is known with certainty only from the adult male.
Barraud (1929) described an allotype female, from Sukna, N. Bengal, which
has apparently been lost. It is not, as stated by Barraud, in the British Muse-
um and there is no record of its ever having been received there. A female
from the Andaman Islands, identified as this species by Barraud, is in the
British Museum and I formerly marked it, provisionally, as a paratype
(Mattingly 1957). Having now had the opportunity to compare it with other
material, I am inclined to doubt that it belongs to the present species. I have,
therefore, omitted it from the following description and noted differences in
the Taxonomic Discussion below. The following description of the female is
based on material from Thailand, provisionally attributed on the basis of a
comparison with males and with Barraud's description of the allotype.
20 Contrib. Amer. Ent. Inst., vol. 5, no. 7, 1970
FEMALE. Head. No pale scales round borders of eyes; eyes well
separated; basal joint of antennal flagellum dark scaled; proboscis about nine-
tenths the length of the fore femur; palpus about one- seventh to one-sixth the
length of the proboscis; clypeus bare. Thovax, Scutal scales with bright gold
reflection shot with red and blue; prescutellar bristles absent; scutellum with
dark scales on all lobes (but see below under Taxonomic Discussion); upper
surface of anterior pronotal lobes covered with silvery scales, narrowly dark
behind; posterior pronotum entirely dark scaled; postspiracular bristles ab-
sent; postnotum with conspicuous tuft of bristles. Wing. Plume scales nar-
row except distally on Ro and, to a less extent, Rg; cell Ro about 2.5 times
the length of its stem or slightly more or less. Legs. Hind femur pale on
about the basal three-fourths to four-fifths of the anterior surface with dark
dorsal stripe reaching to base. Abdomen. Tergum I dark; II-VI with basal
lateral triangular white patches extending ventrolaterally to the posterior edge
of the tergum on II and inwards well towards the mid line; II-V with or without
median pale basal spots which may fuse with the basal lateral pale patches in
some cases; VI and VII with or without a basal pale band; distal sterna dark
apically; sternum II apparently all pale. Terminalia, Postgenital plate
smooth or very slightly emarginate at tip; postatrial plates strongly sclero-
tized; sclerotized portion of spermathecal duct noticeable longer than usual
(? always).
MALE. Head, Antenna with the 2 terminal joints of the flagellum
greatly elongated. Legs. Each fore tarsal claw with a large tooth at base;
one mid tarsal claw with a stout tooth, the other with a very slender, hair-
like tooth at base. Terminalia, Distimere much as in communis but less
expanded apically and with a more sharply marked median fold or ridge; apex
of basimere with a double row of flattened, striated setae; subapical lobe with
a stout, twisted spine and a slender seta immediately adjacent to it; distal
claspette @cl) with appendix of harpago reduced to a small lobe resembling a
larval comb scale; proximal claspette (bcl) with 3 setose lobes, one with very
long, sinuous, flattened setae, their bases fused to form a transparent, stri-
ated sheet; paraprocts with thumb-like appendage at tip; [Xth tergum with
rounded lobes, each with about 4 small setae; I[Xth sternum almost completely
sclerotized with a median lobe bearing about 10 small setae.
PUPA. Not known.
LARVA. Provisionally described from unassociated whole larvae
(see below under Taxonomic Discussion). Head, Antenna moderately long,
curved, lightly spiculate; seta 1-A with 2-4 branches; mouthbrushes with fine,
non-pectinate setae only; seta 5-C with 6-12 branches; 6-C single or with 2
very unequal branches. Abdomen. Comb of 7-14 narrow, more or less uni-
formly fringed scales (cs); siphon about 2.7 - 3.0 times the length of the sad-
dle (3.5 - 4.0 times breadth at base); pecten of 4-8 large teeth @7) mostly with
conspicuous basal denticles, but some of the basal ones sometimes simple;
seta 1-S with 2-3 branches, arising just beyond distal pecten tooth, longer than
diameter of siphon at point of attachment; saddle with distal edge spiculate e~-
specially around bases of caudal setae; seta 1-X single or bifid; 2-X with 2-3
branches; 3-X single; anal papillae bluntly rounded, the dorsal pair longer than
Mattingly: Heizmannia in Southeast Asia 21
the ventral; the dorsal pair about 3 times as long as the saddle in specimens
from Chiang Mai (much shorter than this in specimens from Trad and Rangoon
but these appear to be immature); ventral brush with 6 setae in the barred
area and 1-2 smaller precratal setae; all these setae single or bifid except for
the most proximal seta in the barred area which is trifid in one Specimen.
TYPE DATA. Lectotype male on pin, with terminalia on celluloid
mount, in British Museum; type locality: Andaman Islands (see Mattingly
1957). Allotype female believed lost; type locality: Sukna, North Bengal,
INDIA.
DISTRIBUTION. Specimens examined: INDIA, North Bengal, Sukna
1 male. Andaman Islands, 2males. BURMA, Rangoon 2 whole larvae.
THAILAND, Chiang Mai, Doi Suthep 1 female, 1 male, Ban Pha Tak 1 female,
Ban Kee Lek 1 female, Huey Chang Kien 1 female; Tvat, Ko Chang 3 females,
1 whole larva; Unnamed localities 1 female, 2 whole larvae. Other Distribu-
tion. Recorded by Causey (1937) from locality between Bon Pin and Chieng
Moeang (? = Ban Chiang Muang) THAILAND. A record of "H. communis"
from Ceatigs CHINA (Borel 1930) may also refer to the present species (see
below). |
TAXONOMIC DISCUSSION. Barraud (1929, 1934) mentions the pres-
ence of silvery scales at the apex of the mid lobe of the scutellum. No such
scales are present in the Andaman specimens in the British Museum or in any
of the specimens from Thailand. They may have been present in the missing
allotype. No associated larval or pupal skins are available but some whole
larvae from Thailand resemble those from Burma attributed by Barraud to the
present species. No other Burmese material has been preserved but the pres-
ence of covelli in Thailand has been confirmed from male terminalia. The de-
scription of the female adult and, particularly, the larva of "H.communis"’,
from Cochin China, by Borel (1930) would agree better with the present spe-
cies. Females seem most likely to be confused with those of scintillans and,
perhaps, mattinglyi. Thurman (1959) treats the color of the scutal scaling as
diagnostic but this is based on a misinterpretation of Barraud's description,
which is misleading. She does not appear to have seen any specimens.
BIOLOGY. Not clearly recorded as attacking man. Causey (1937)
records it as biting by day in jungle but says it was collected from a horse.
The whole larva described here came from tree holes and a coconut. The
paratype male from the Andamans is labeled as from a bamboo stump and the
Thailand male and the male from Sukna as from tree holes. Records by Jolly
(1933) from a variety of artificial containers, as well as from tree holes and
bamboos, in Rangoon require confirmation.
HEIZMANNIA (HEIZMANNIA) DEMEILLONI n. sp.
(Figs. 17,9; 18,2 terminalia, pupa, claws)
Closely related to himalayensis Edwards, of which it may be a sub-
species. It is, however, distinguished from this, and all other species in the
genus, by the striking scutal ornamentation. H. himalayensis has not been
22 Contrib. Amer. Ent. Inst., vol. 5, no. 7, 1970
recorded from Southeast Asia but it occurs in North Bengal and may well be
found in Kast Pakistan. Some notes on this species will be found in the Taxo-
nomic Discussion below.
FEMALE. Head, No continuous border of white scales round eye
margins although the usual ventrolateral patches of white scales extend in-
wards, narrowly, well towards the median patch between the eyes; eyes wide-
ly separated as far forwards as the tori; basal joint of antennal flagellum with
dark scales only; proboscis slender, about 1.1 times the length of the fore
femur; palpus about one-tenth the length of the proboscis; clypeus bare.
Thorax. Scutal scales narrower than in other species (except himalayensis)
with bright, matt gold reflection, quite different from the brassy reflection
seen in covelli and scintillans and with much less tendency to the formation of
iridescent colors than in those species; prescutellar bristles absent; mid lobe
of scutellum with broad silvery scales and some dark ones, lateral lobes dark
scaled; upper surface of anterior pronotal lobes almost entirely covered with
broad white scales; posterior pronotum with similar scales below; some dark
scales above; postspiracular bristles absent; postnotum with pale colored
bristles, without scales. Wing. Plume scales broad; cell Rog variable but
long in all cases, from about 3.0 - 4.5 times the length of its stem. Legs,
Hind femur pale on basal four-fifths or more of anterior surface with dark
dorsal stripe broadening towards tip; this stripe very narrow on basal half but
reaching to base; all tarsal claws simple. Abdomen. Tergum I with a small
pale spot; laterotergite pale scaled; terga II-IV dark above; V with inner edges
of basal lateral pale spots visible on dorsal surface; VI and VII with basal pale
bands; all terga with triangular basal lateral pale spots, prolonged along ven-
trolateral edge to posterior edge of terga II and III; sterna with dark apical
bands (II apparently all pale). Terminalia. Postgenital plate square-ended
but without apical emargination.
MALE. Not known.
PUPA. Cephalothorax. Seta 6-C longer than usual, not very much
shorter than 7-C, single; seta 7-C somewhat longer, more slender, single or
bifid. Abdomen. Seta 1-II (preserved in only one specimen) with 3-4 primary
branches, little or no secondary branching; 5-II small and very delicate; 9-II-
VI minute, very inconspicuous; 9-VII single; 9-VUI with 3-4 branches. Paddle.
About 1.5 - 1.7 times as long as its greatest breadth; fringe of fine hairs
shorter and scantier than in most species; seta 1-P broken.
LARVA. The slide preparation of the larval skin was lost before it
could be illustrated. Head, Antenna long, curved, moderately stout, spicu-
late; seta 1-A bifid, arising unusually near tip; mouthbrushes with fine, non-
pectinate hairs only; setae 5-C and 6-C not seen. Abdomen. Comb an irreg-
ular patch of 9-12 spines with fine basal fringe and strongly hypertrophied
median denticle; siphon about 3 times the length of the saddle (about 3.5 times
the breadth at base); pecten of 4-7 spines, all with strongly developed basal
denticles; seta 1-S arising just beyond terminal pecten tooth, bifid, about e-
qual in length to the breadth of the siphon at point of attachment; saddle with
apex lightly spiculate; seta 1-X missing; 2-X with 4 branches; 3-X single; anal
Mattingly: Heizmannia. in Southeast Asia 23
papillae about 2 times as long as saddle, relatively narrow, cylindrical,
bluntly pointed, the ventral pair slightly shorter than the dorsal; ventral
brush apparently with 8 setae, all of them missing except 1 of the 2 precratal
setae which is bifid with one branch split secondarily into 2.
TYPE DATA. Holotype female, on pin, with associated pupal skin
on Slide in U. S. National Museum; type locality: Kalaw, BURMA.
DISTRIBUTION. Specimens examined: BURMA, Kalaw 3 females,
1 larval and 3 pupal skins. THAILAND, Chiang Mai, Doi Suthep 1 female.
TAXONOMIC DISCUSSION. As noted above, himalayensis is closely
related to the present species. It differs chiefly in the coloration of the scutal
scaling which has a characteristic dark blue reflection. Other differences are
as follows: proboscis slightly shorter, about equal in length to fore femur;
cell Rg about 2.5 - 3.5 times the length of its stem; plume scales narrower
(but the specimen of demeilloni from Thailand has them narrower than in the
Burmese specimens); hind femur with dark dorsal stripe not reaching base;
white basal lateral patches on abdominal terga narrower, prolonged back-
wards, if at all, only on tergum II; pale basal median spots or bands some-
times present on II-V; postgenital plate slightly more rounded than in demeil-
loni; terminalia otherwise very Similar. Early stages very similar except
that the larva of himalayensis has a much shorter antenna,
BIOLOGY. The Burmese specimens were bred out from bamboo
stumps,
HEIZ MANNIA (HEIZMANNIA) FUNEREA. (LEICESTER)
(Figs. 19, 2; 20, °° terminalia, “ claws; 16, larva)
Wyeomyia funerea Leicester 1908, Cul. Malaya, 252 (9).
Closely resembles indica and scintillans, differing in the greatly re-
duced pale scaling on the prothoracic lobes. The males are easily recognized
by the plumose mid tarsi which are unique in the genus.
FEMALE. Head. Top of head dark with ventrolateral pale patches
and small pale spot between the eyes; basal joint of antennal flagellum with
pale scales, dark in some lights; proboscis about 1.1 times as long as fore
femur; clypeus bare; palpus about one-fifth of the length of the proboscis.
Thorax. Scutal scales with green, purple and blue reflections; scutellum
dark scaled; anterior pronotal lobes largely dark scaled with small pale spot
on anterior border; posterior pronotum dark scaled; postspiracular bristles
absent; postnotum with tuft of dark setulae. Wing. "Plume scales mostly nar-
row, broader on veins Rg and Rg; cell Rog about 1.8 - 3.0 times as long as its
stem. Legs. Hind femur with about the basal two-thirds to three-fourths of
the anterior surface pale, the dark apical band prolonged ventrally to base.
Abdomen. Tergum I dark; laterotergite white scaled; terga II-VI dark above
with triangular basal lateral pale patches prolonged inwards well towards mid
line but not forming a complete basal band on any segment; basal lateral pale
24 Contrib. Amer. Ent. Inst., vol. 5, no. 7, 1970
spot on tergum II prolonged along the ventral border to the posterior edge of
the tergum; the corresponding spot only slightly prolonged on other terga;
distal sterna with dark apical bands; sternum II apparently entirely pale;
sternum VII pale at base in some specimens, entirely dark in others. Termi-
nalia, Postgenital plate expanded at apex.
MALE. Head, Antenna lightly plumose; the two terminal joints
greatly elongated. Legs. Fourth and fifth mid tarsal segments with tufts of
sub-erect scales; hind tarsus with the last two segments missing; other seg-
ments without any such scales; one fore and one mid tarsal claw with slender
tooth at base. Terminalia, Much as in scintillans but with leaflike setae at
apex of basimere somewhat broader, 2 spines on subapical lobe and distal
claspette (cl) quite different, more as in persimilis, propinqua and proxima
n. Spp.
PUPA. Not Known.
LARVA. Head. Antenna long, slender, lightly spiculate; seta 1-A
with 4-5 branches, arising at or slightly before half way; mouthbrushes with-
out pectinate setae; seta 5-C with about 10-15 branches; 6-C with 2very un-
equal branches. Abdomen. Comb a patch of about 13-20 scales (cs), with
uniform apical fringe, and spines with hypertrophied median denticle; siphon
about 3 times the length of the saddle (about 3.5 times its breadth at base);
pecten of 6-9 slender teeth (p/) without secondary denticles; seta 1-S missing
from both the available specimens, its base situated just beyond the distal
pecten tooth; saddle with apex lightly spiculate; seta 1-X long, Single; 2-X
bifid; 3-X single; anal papillae short, broad, bluntly rounded at tip, subequal;
ventral brush with 6 setae in barred area, all single or bifid, and 2 precratal
setae, both single.
TYPE DATA. Holotype female on pin in British Museum; type local-
ity: Jungle 6 miles from Kuala Lumpur on Pahang Road, Selangor, WEST
MALAYSIA.
DISTRIBUTION. WEST MALAYSIA, Selangor, 6th mile, Pahang
Road 1 female, 10th mile, Banting Road 1 female, Ampang Forest Reserve
14 females, 1 male. EAST MALAYSIA, Sabah, Tawau 2 females. SINGA-
PORE 1 female, 2 unassociated larval skins, 4 immature whole larvae.
TAXONOMIC DISCUSSION. Though closely related to scintillans and
formerly placed in the synonymy of that species (Mattingly 1957) it is clear
from the secondary characters of the male, described here for the first time,
that this is a distinct species. Two specimens from Singapore placed by
Edwards under funerea included a male of indica, I therefore tentatively
suggested that the larva described by Edwards and Given (1928) as funerea
might, in fact, be that of indica, Comparing the other specimen from Singa-
pore, a female, with the new material of funerea from Ampang, however, I
am satisfied that it is the present species. It seems best, therefore, to ac-
cept Edwards and Given's identification provisionally and to treat the larva of
indica as still unknown.
BIOLOGY. Edwards and Given's larvae came from a tree hole.
Mattingly: Heizmannia in Southeast Asia 25
HEIZMANNIA (HEIZMANNIA) INDICA (THEOBALD)
(Figs. 21, ?; 22, °9 terminalia, « claws)
Phoniomyia indica Theobald 1905, Ann. hist.-nat. Mus. hung. 3: 115 (o*).
Wyeomyia metallica Leicester 1908, Cul. Malaya, 251 (?). NEW SYNONYMY
Closely resembles scintillans and funerea but is distinguishable by
the more uniformly blue color of the scutal scaling and the truncation of the
basal lateral pale triangles on the abdominal terga. The advent of further
material has strengthened the view that Leicester's "9, indica" was in fact
veidi (Mattingly 1957). It has also made it possible to synonymize his
metallica with the present species with some assurance. The Indian species
attributed to metallica by Edwards (1922) and Barraud (1929, 1934) is evi-
dently distinct. It will require to be redescribed under a new name. It dif-
fers markedly in the male terminalia and has decidedly narrower wing scales,
especially on the veins of the upper fork.
FEMALE. Head. Eyes well separated; ventrolateral patches of pale
scales on head extending in towards mid line but eye margins without continu-
ous border of pale scales; basal joint of antennal flagellum with a small spot
of pale scales and, usually, some dark ones; proboscis long and slender,
about 1. 2 times as long as front femur; palpus about one-fifth or less the
length of the proboscis; clypeus bare. Thovax. Scutal scales mainly with
dark blue and blue-green reflections; no prescutellar bristles; scutellar scales
all dark; anterior pronotal lobes with pale scaling variable in extent but al-
ways limited to the anterior edge, dark scales posteriorly; posterior pronotum
wholly dark scaled; postspiracular bristles absent; postnotum with a strongly
developed group of dark bristles. Wing. Plume scales on Ro and Rg narrow
to moderately broad; cell Ro from less than 2 times to about 2. 5 times as
long as its stem. Legs. Hind femur pale on about the basal two-thirds to
seven-tenths with broad, dark dorsal stripe continued to base and also with a
dark ventral stripe, variously developed (much as in scintillans, Figure 39,
p. 39). Abdomen. TergaI-VI dark above; VII sometimes with a narrow pale
basal band or small basal lateral patches, sometimes all dark; II-VI with
basal lateral white spots truncated, not extending far inwards or tapering to a
point towards mid line, extending posteriorly along ventrolateral edge only on
II and to some extent on III; sterna with broad dark apical bands. Tevminalia,
Cerci short and broad; postgenital plate bluntly rounded at apex.
MALE. Head, Antenna with the 2 terminal joints greatly elongated.
Legs. One fore tarsal claw with a slender, straight tooth at base; the other
simple; mid tarsal claws similar but the single tooth even more slender.
Terminalia. Distimere resembling that of covelli but differing in various de-
tails; apex of basimere with a row of 6 short, narrow, striated setae; subapi-
cal lobe with a single large twisted spine; distal claspette (cl) with 2 groups
of stout setae, one group with hooked tips; no harpago; proximal claspette
(cl) with 2 groups of flattened setae, the longer group with fimbriated tips,
and a broad appendage recalling the fused basal portions of the long setae in
25°. Contrib. Amer. Ent. Inst., vol. 5, no. 7, 1970
chengi and covelli but with these setae reduced to short, blunt, transparent,
faintly striated leaflets; paraprocts with thickened, doubly recurved tips; [Xth
tergum strongly sclerotized; each lobe with about 6-8 fine hairs; [Xth sternum
almost completely sclerotized, with a stout bristle near apex.
PUPA and LARVA. Not known.
TYPE DATA. 4d. indica (Theobald), holotype male on pin in Hungar-
ian National Museum; type locality: SINGAPORE. 4H. meiallica (Leicester),
holotype female on pin in British Museum; type locality: Bukit Kutu, Selangor,
WEST MALAYSIA. |
DISTRIBUTION. Specimens examined: SINGAPORE 2 males. WEST
MALAYSIA, Selangor, Bukit Kutu 1 female, Pahang Road 1 female, Ampang
1 female, Ulu Gombak 12 females, 1 male, Banting Road 1 female; Pevak, Bt.
Pulau Tiga 1 female; Pahang, Kuala Tahan 1 male, Kuantan 1 female. Other
Distribution. The record by Causey (1937) from THAILAND ("between Bon
Pin and Chieng Moeang"') may refer to veidi (see p. 35).
TAXONOMIC DISCUSSION. Additional characters which may be of
diagnostic value are the small size of the present species (wing length at most
2.5 mm. in specimens assignable with confidence), mainly bluish scutal
scaling, all dark first hind tarsus and generally pale scutal integument. The
larva at one time attributed provisionally to this species is now assigned to
funerea (see p. 23 ). The early stages of the present species remain unknown.
Records of "H. indica” from India (Barraud 1929, 1934, Edwards 1922), and
— from Malaya (Leicester 1908) are considered to refer to reidi (see p.
35).
BIOLOGY. The holotype of metallica was taken in a house. Other
adults have been taken on man in forest in daytime.
HEIZMANNIA (HEIZMANNIA) MACDONALDI MATTINGLY
(Figs. 23,9; 24,0 terminalia, claws; 25, pupa, larva)
Heizmannia macdonaldi Mattingly 1957, Cul. Mosq. Indomalay. Area 2: 47
(o*, oF, P*, 1*),
Heizmannia nivirostris Lien 1968, Trop. Med., Nagasaki, 10: 139 (o*, 9*, P*,
L*), NEW SYNONYMY.
_ This species is very similar to communis, the 2 evidently constituting
a closely related sibling species pair. The females differ constantly in the
scaling of the posterior pronotum (dark in communis, pale in the present spe-
cies) and there are small, but apparently constant, differences in the male
terminalia. The larvae and pupae cannot be separated in every case but, on
present evidence, the overlap is only slight.
FEMALE. Head. Top of head dark with lateroventral white spots
and a large white spot between the eyes which are widely separated; eye mar-
gins largely, usually wholly, dark; basal joint of antennal flagellum with whit-
ish or pale brown scales, appearing dark in some lights; proboscis approxi-
mately equal in length to fore femur, rather more thickened towards tip than
Mattingly: Heizmannia in Southeast Asia 27
in most species, with a distinct pale line below towards base; palpus from
one-fifth to one-fourth of the length of the proboscis; clypeus bare. Thovax.
Scutal scales with subdued, dull bronze reflection; prescutellar bristles ab-
sent; scutellar scales all dark; anterior pronotal lobes wholly or largely pale
scaled; posterior pronotal scales mostly whitish or pale brown; postspiracular
bristles absent; postnotal bristles dark. Wing. Plume scales broad; cell R
about 3.0 - 3.5 times as long as its stem. Legs. Hind femur pale below on
anterior surface for about the basal six-sevenths, more narrowly so towards
tip. Abdomen, Tergum I dark; terga II-VI with basal lateral pale patches
prolonged backwards along ventral border on II and to a variable extent, onIII;
V-VII dark above or, occasionally, with pale basal bands; sterna with dark
apical bands. Terminalia, Postgenital plate emarginate at apex, very much
as in communis.
MALE. Head, Antenna of normal type; proboscis much as in female.
Legs. One fore tarsal claw with a long, slender tooth at base; the other sim-
ple. Mid tarsal claws both simple. Terminalia. Very like those of commu-
nis, differing mainly in the slender paraprocts and the narrower 'neck' at the
base of the appendage of the harpago (i). Small differences in the ornamenta-
tion of the proximal claspette (pcl) do not seem to be constant.
PUPA. Cephalothorax, Seta 6-C single, shorter than 7-C which is
single to tetrafid. Abdomen. Seta 1-II strongly developed, semi-palmate;
o-II short, very slender, single; 9-II-VI small, delicate; 5-IV and 5-V about
equal in length to the following segment or somewhat longer or shorter; 5-VI
about half as long as the following segment or less; 9-VII and 9-VIII single to
tetrafid and single to pentafid respectively. Paddle, From about 1.6 - 2.0
times as long as its greatest breadth; seta 1-P single to trifid; male genital
sac with apical horns poorly differentiated.
LARVA. Head, Antenna dimorphic, as in communis, shorter in
those larvae with pectinate mouthbrush setae, longer in those with the filter-
feeding type of mouthbrush, spiculate for most of its length; seta 1-A with 2-6
branches; mouthbrushes with simple hairs in 15/16 larvae examined, with
pectinate setae in the remaining one; seta 5-C with 5-11 branches; 6-C usually
with 3-5 branches (see under Taxonomic Discussion). Abdomen. Comba
mixture of about 13-23 scales (cs), with uniformly fringed apex, and spines
with hypertrophied median denticle, the former predominating; siphon about
3.5 - 5.0 times the length of the saddle (about 5-8 times its breadth at base);
pecten of 4-12 slender spines (pi), usually with the bases less expanded and
the secondary denticles smaller than in communis; seta 1-S bifid; arising well
beyond the distal pecten tooth, somewhat longer than the diameter of the siphon
at point of attachment; saddle with apex lightly spiculate; seta 1-X usually bi-
fid or trifid, sometimes longer and single; 2-X with 2-4 branches; 3-X single;
anal papillae large, cylindrical, bluntly rounded at tip, the upper pair much
longer than the lower; ventral brush with 6 setae in the barred area, 1-4 of
these bifid (or one, rarely, trifid), the others single, and with 2 smaller setae
arising outside the barred area.
28 Contrib. Amer. Ent. Inst., vol. 5, no. 7, 1970
TYPE DATA. %. macdonaldi Mattingly, holotype male and allotype
female, on pins, with associated larval and pupal skins, in British Museum;
type locality: Ulu Langat Forest Reserve, Selangor, WEST MALAYSIA. 4,
nivirostris Lien, holotype male and allotype female, with associated larval
and pupal skins, in Provincial Malaria Research Institute, Taipei, Taiwan;
type locality: Laitou, Fanlu, Chiai, TAIWAN.
DISTRIBUTION. Specimens examined: THAILAND, Tak, Doi Sam
Sao 1 female, 1 larval skin, 1 pupal skin; Ranong, Kra Buri 1 male, 1 pupal
skin; Nakhon Si Thammarat, Ban Thuan Lek 1 whole larva. WEST MALAYSIA,
Selangor, Ulu Gombak 1 female, 1 larval skin, 1 pupal skin, Ulu Langat 7 fe-
males, 14 males, 12 larval skins, 12 pupal skins. TAIWAN, Chiai, Laitou
1 paratype female of nivirostris, 1 whole pupa with associated larval skin of
same. Other Distribution. Recorded in TAIWAN only from the type locality
of nivirostris.
TAXONOMIC DISCUSSION. Distinctions from communis have been
discussed under that species. All identifiable larvae from the mainland seen
by me have seta 6-C with 3 or more branches at least on one side. However,
the only Taiwan larva available to me has this seta bifid on both sides and
with one branch markedly smaller than the other. Distinction from communis,
and even from scintillans and funerea, may not therefore always be possible.
The female from Taiwan seen by me has a pale basal band on abdominal terga
V-VI. In mainland specimens I have seen such a band on tergum VII only.
This, however, is a variable character in so many species that I do not attach
very much importance to it. None of the characters given by Lien for distin-
guishing nivirostris from the present species are reliable. All fall within the
range of variation of my mainland material. I have not seen any males from
Taiwan and Lien's figure of the terminalia shows no details of the ornamenta-
tion of the proximal claspette. So far as I can tell, however, there is no sig-
nificant difference from the present species. I have therefore relegated
nivivostris to the synonymy.
BIOLOGY. Breeding places include split and bored bamboos
(Macdonald & Traub 1960) and a bamboo stump at 1000 m. (Lien 1968). The
larva from Ban Thuan Lek came from a tree hole. Macdonald & Traub note
a preference for dead or dying bamboos.
HEIZMANNIA (HEIZMANNIA) MATTINGLYI THURMAN
(Figs. 24, 9 terminalia; 26, 2)
Heizmannia mattinglyi Thurman 1959, Univ. Maryland Agric. Exp. Sta. Bull.
A-100: 70 (9).
Known only from the adult female. Apparently distinguished from all
other species of the genus by the presence of pale scales on the lateral, as
well as the median, lobes of the scutellum. It closely resembles covelli in
other respects.
Mattingly: Heizmannia in Southeast Asia 29
FEMALE. Head. Top of head dark except for the usual ventrolater-
al white patches and white spot between the eyes; eyes well separated; basal
joint of antennal flagellum with dark scales; proboscis about equal in length to
fore femur; palpus about one- seventh of the length of the proboscis; clypeus
bare. Thorax. Scutal scales mainly bright gold with greenish reflections;
prescutellar bristles absent; scutellum with white scales on all lobes and some
dark scales intermixed; anterior pronotal lobes almost entirely pale scaled
with some dark scales posteriorly; posterior pronotum dark scaled; postspira-
cular bristles absent; postnotum with a tuft of dark bristles. Wing. Plume
scales narrow, broader on Rj and Rg; cell Rg about 3.0 - 3.5 times as long
as its stem. Legs. Hind femur pale on about the basal four-fifths with dark
dorsal stripe not quite reaching base. Abdomen. Tergum I dark; II-VI with
basal lateral pale patches prolonged posteriorly along lateral edges on II and
to some extent on III; segments II-VI otherwise dark or with more or less
complete pale basal bands; VIL with a complete pale basal band in all cases;
sterna II and III apparently all pale; posterior sterna with apical dark bands.
Terminalia. Apparently indistinguishable from those of covelli.
MALE, PUPA and LARVA. Not known.
TYPE DATA. Holotype female, on pin, with terminalia on slide, in
U. S. National Museum; type locality: Rong Kwang, Phrae, THAILAND.
DISTRIBUTION. Specimens examined: THAILAND, Phrae, Rong
Kwang 2 females; Chiang Mai, Prao 1 female, Chiang Mai 1 female, Doi
Suthep 2 females, Chiang Dao 1 female; Udon Thani, unnamed locality 1 fe-
male; Nan, Ban Salik 1 female. CAMBODIA, Sre Klong | female, Prey
Plidan 1 female. Other Distribution. The U. S. National Museum has a fe-
male, from Ban Pha Tak, Chiang Mai, THAILAND, attributed to this species
but now fragmentary and no longer identifiable. Records other than from the
type locality require confirmation.
TAXONOMIC DISCUSSION. The paratype from Rong Kwang has the
abdomen missing. The remaining specimens differ from the holotype in the
presence of conspicuous pale basal bands on abdominal terga II-VI. However,
this is a variable character in some other species, notably covelli, and its
taxonomic significance is doubtful. The holotype and paratype appear to have
the plume scales narrower than in the other specimens but both have the wings
badly rubbed. The only reliable distinction from covelliseems to be the pres-
ence of pale scales on the lateral lobes of the scutellum. The significance of
this character cannot be properly assessed until associated material is avail-
able.
BIOLOGY. Adults were taken attacking man in jungle and bamboo
forest and, in the case of the holotype and paratype, in partial shade on a
hillside.
30 Contrib. Amer. Ent. Inst., vol. 5, no. 7, 1970
HEIZMANNIA (HEIZMANNIA) PERSIMILIS n. sp.
(Figs. 27, 9; 28, 2 terminalia, “ claws; 29, pupa, larva)
This is one of several species which may previously have been re-
corded as gveenii. The others are chengi, propinqua n. Sp.,proxima n. sp.
and scanloni n. sp. All of them have the posterior pronotum pale scaled and
very narrow wing scales, a combination of characters previously considered
diagnostic for greenit.
FEMALE. Head, Top of head dark except for pale ventrolateral
patches and pale spot between eyes; basal joint of antennal flagellum with pale
scales; proboscis relatively stout, about 1.1 times as long as front femur;
clypeus bare, palpus about one-tenth to one-eighth of the length of the probos-
cis. Thorax. Scutal scales with metallic, mainly green reflection, blue in
some lights; a small spot of broad, white scales on lateral edge, immediately
in front of wing root; prescutellar bristles absent; scutellum entirely dark
Scaled; anterior pronotal lobes with a border of pale scales round the entire
anterior and outer edge; posterior pronotum entirely pale scaled; postspiracu-
lar bristles absent; postnotum with a conspicuous tuft of moderately pale bris-
tles, Wing. Plume scales all very narrow; cell Rj about 2.3 - 2.8 times as
long as its stem. Legs. Hind femur pale on about the basal four-fifths; all
tarsal claws simple. Abdomen. TergaI-IV all dark above or IV with a small
median basal pale spot; remaining terga dark above or with more or less com-
plete, narrow pale basal bands; tergum II with squarish basal lateral pale.
patches extending back to the posterior border; III-VI with triangular basal
lateral pale patches tending to be prolonged posteriorly on anterior, and in-
wards towards center on posterior segments; sterna III-VII with broad, dark
apical bands. Tervminalia, Postgenital plate with rounded apex; postatrial
plates strongly sclerotized.
MALE. Head, Antenna with the 2 terminal joints elongated as usual.
Legs. One fore tarsal claw with a tooth arising near base; the other simple;
both mid tarsal claws simple. Terminalia. Distimere without special fea-
tures, very much as in proxima n. Sp.; apex of basimere with a single row of
narrow, Slightly flattened setae; subapical lobe with a single spine; distal
claspette dcl) with 2 pilose lobes, without harpago, very much as in propinqua
n. Sp. and proxima n. sp.; proximal claspette (bcl) with numerous setae; a
few of which are stouter than the rest while others are greatly elongated; para-
procts with 1 large and 1-2 smaller teeth at apex; I[Xth tergum with lobes
closely approximated, each with 4-6 delicate hairs; [Xth sternum completely
sclerotized with 1-2 stout bristles at apex.
PUPA. Cephalothorax, Seta 6-C single, shorter than 7-C which is
single or bifid. Abdomen. Seta 1-II relatively conspicuous, with several
primary branches, one or more of which may show secondary branching; 5-II
very Slender, single or bifid; 9-II-VI minute; 5-IV variable, usually about as
long as the following segment, occasionally only half as long; 5-V longer than
the following segment; 5-VI about as long as the following segment or a little
Shorter; 9-VII single; 9-VIII bifid or trifid. Paddle. About 1.6 - 2.3 times
as long as its greatest breadth; fringe tending to be less extensive than in most
Mattingly: Hezzmannia in Southeast Asia 31
Species, with fine hairs and a few spicules inside these; seta 1-P single; male
genital sac with well marked horns. |
LARVA. Head. Antenna variable in length, curved, spiculate; seta
1-A arising shortly before half way in those larvae with short, and slightly
beyond half way in those larvae with long, antennae, with 3-4 branches; mouth-
brushes without pectinate setae; seta 5-C with 6-13 branches; 6-C single or,
more usually, with 2 very unequal branches. Abdomen. Comb with 16-28
teeth (cs), comprising mixed scales, with more or less uniform fringe, and
Spines with hypertrophied median denticle; siphon about 3.0 - 3.7 times the
length of the saddle beet 3.7- 4.5 times its breadth at base); pecten of 5-10
teeth (p/), some simple, others with well developed basal denticles; seta 1-S
with 2-3 branches, longer than diameter of siphon at point of attachment, a-
rising just beyond terminal pecten tooth; saddle with apex strongly spiculate;
seta 1-X single; 2-X single to trifid, usually bifid; 3-X single; anal papillae
long, bluntly rounded at tip, narrower than in most species; ventral brush
with 6 setae in the barred area and 1-2 precratal setae, all single or bifid.
TYPE DATA. Holotype male, on pin, with terminalia and associated
larval and pupal skins on slides, in British Museum; type locality: 15th mile,
Ulu Gombak, Selangor, WEST MALAYSIA,
DISTRIBUTION. Specimens examined: THAILAND, Tak, Khao Salak
Phra 7 females, 6 males, 6 larval skins, 13 pupal skins. WEST MALAYSIA,
Selangor, Ulu Gombak 3 females, 2 males, 2 larval skins, 2 pupal skins.
Other Distribution, The following unassociated specimens are provisionally
attributed to the present species, though excluded from the paratype series.
THAILAND, Chiang Rai, Doi Sam Sao 3 whole larvae; Ranong, Ko Chang Is.
1 larval skin, 1 pupal skin; Nakhon Nayok, Pha Klory Mai 1 pupal skin. WEST
MALAYSIA, Selangor, Ulu Gombak 1 female. VIETNAM, An Tuc 1 female,
Cam Ranh 26 females.
TAXONOMIC DISCUSSION. Specimens from Vietnam, tentatively
placed under this species, by reason of the pale scaled posterior pronotum,
have the wing scales rather broader than in topotypic persimilis n. sp. They
also show a tendency to prolongation of the apical dark area along the under-
Side of the hind femur, approaching scintillans in both respects. They can at
present only be provisionally ascribed. As noted above, this species may
formerly have been confused with greenii, It seems, however, to be more
closely related to scintillans, as also do propingua n. sp. and proxima n. sp.
BIOLOGY. The paratypes from Khao Salak Phra were bred out from
a tree hole in primary rain forest at 2700 ft. The whole larvae from Doi Sam
Sao and the pupa from Ko Chang Is., provisionally attributed to the present
Species, were found in holes in tree stumps in forest. Some Malayan adults
were bred out from a tree hole about 6 ft. above ground, containing 'peaty'
water. Others were taken on man,
32 Contrib. Amer. Ent. Inst., vol. 5, no. 7, 1970
HEIZMANNIA (HEIZMANNIA) PROPINQUA n, sp.
(Figs. 30, 9;31,% terminalia, claws; 32, pupa, larva)
Closely resembles persimilis n. sp. and proximan. sp. but differs
from both in the ornamentation of the apex of the male basimere and proximal
claspette.
FEMALE. Head, Top of head with pale scales largely confined to
the median and ventrolateral spots but with a few pale-tipped scales round eye
margins; basal joint of antennal flagellum with pale scales; proboscis relative-
ly stout, approximately equal in length to fore femur; clypeus bare; palpus
about one-sixth of the length of the proboscis. Thovax. Scutal scales with
rather dull, mainly green reflection but distinctly metallic (in contrast to
complex), much as in persimilis n. sp.; prescutellar bristles absent; alllobes
of scutellum dark scaled, upper surface of anterior pronotal lobes dark except
for a rim of whitish scales round the anterior border; posterior pronotum en-
tirely pale scaled; pleuron with the usual ornamentation of broad, white scales
postspiracular bristles absent; postnotum with a small tuft of pale bristles.
Wing. All plume scales narrow; cell Ro about 2.5 times as long as its stem.
Legs. Anterior surface of hind femur With a broad creamy stripe on about
the basal five-sixths; dark dorsal stripe continued to base. Abdomen, Terga
I-VI nearly all dark above; II-VI with large, creamy basal lateral triangles,
prolonged backwards along the ventrolateral edge on II-V; VII and rest of ab-
domen missing; sterna with dark apical bands except, apparently, onIl. Ter-
minalia. Missing.
MALE. Head, Antenna with the 2 terminal joints elongated as usual.
Legs. One fore tarsal claw with a long, slender tooth at base; the other sim-
ple; both mid tarsal claws simple. Terminalia. Distimere with part ofouter
edge folded, producing a small spur; apex of basimere with 2 rows of slightly
flattened setae, the proximal row longer; subapical lobe single with a long,
stout, sinuous spine; proximal claspette @cl) with a tuft of relatively short,
flattened setae, several of which are twisted and darker than the remainder,
and with about 5 longer setae, with leaf-like distal expansions, apparently
fused at base, arising from a small apical projection; distal claspette dcl)
with 2 pilose lobes devoid of ornamentation; paraprocts with 1 large and 1
small tooth at apex; [Xth tergum as figured; [Xth sternum uniformly sclero-
tized, without median lobe. 7
PUPA. Cephalothorvax, Seta 6-C shorter than 7-C, bifid; 7-C bifid
on one side, single on the other. Abdomen, Seta 1-II with 8 very delicate
branches; 5-II small, delicate, single; 9-II-VI minute; 5-IV-VI single, about
equal in length to the following segments on IV and VI, about half as long again
on V; 9-VII single; 9- VIII bifid on one side, trifid on the other. Paddle. About
1.9 times as long as its greatest breadth; fringe relatively short and delicate;
seta 1-P single.
LARVA. Head, Antenna moderately long, curved, spiculate; seta
1-A with 2-3 branches, arising well before half way; inner mouthbrush setae
relatively slender but pectinate; seta 5-C with 5-6 branches; 6-C with 2
branches, one much stouter than the other. Abdomen, Comb a patch of 20-24
Mattingly: Heizmannia in Southeast Asia 33
teeth (cs), some with hypertrophied median denticle, others with uniform
fringe, the denticles in all cases relatively coarse; siphon about 3.5 times as
long as saddle (about 4.5 times its breadth at base); pecten with 6-9 teeth @i),
all with secondary denticles which are most conspicuous on the more distal
teeth; seta 1-S bifid, with one branch split distally, longer than the diameter
of the siphon at point of attachment, arising just beyond the distal pecten tooth;
saddle with upper portion of distal edge strongly spiculate; seta 1-X missing;
2-X bifid; 3-X single; anal papillae broad, cylindrical, bluntly rounded at tip;
the lower pair slightly longer than the upper; ventral brush with 6 setae in the
barred area and 2 smaller precratal setae; none of these with more than 2
branches,
TYPE DATA. Holotype male and allotype female on pins in British
Museum, the latter with associated larval and pupal skins; type locality: Ulu
Gombak, Selangor, WEST MALAYSIA.
DISTRIBUTION. Specimens examined: WEST MALAYSIA, Selangor,
Ulu Gombak 1 female with associated larval and pupal skin, 1 male. Other
Distribution. A whole larva from Ban Pha Man, THAILAND, in the U. S.
National Museum, agrees with the above description and may belong to the
present species but the record requires confirmation. A female from Ko
Chang, Ranong, THAILAND, also in the U. 8. National Museum, similarly
agrees but could belong to scanloni n. sp. the female of which is unknown.
(See under Taxonomic Discussion, below).
TAXONOMIC DISCUSSION. As already noted (p. 30), this is one ofa -
group of species any or all of which may previously have been confused with
greenit. Among these its closests affinities, to judge from the male termi-
nalia, would be with persimilis n. sp. and proximan. sp. The whole larva
from Ban Pha Man, mentioned above, keys to the present species but differs
as follows: seta 5-C with 7-8 branches; spines at bases of pleural setae
slightly larger; comb with 10-12 teeth; siphon about 3.2 times as long as sad-
dle; pecten with 5-6 teeth; seta 1-S bifid; distal edge of saddle very lightly
Spiculate; seta 1-X bifid; anal papillae missing. In view of the fact that sever-
al of its relatives are so imperfectly known it seems desirable to query re-
cords from Thailand until these can be supported with males.
BIOLOGY. The paratype female was bred out from a tree hole, about
3 ft. above the ground, containing clean water and sediment. The larva from
Ban Pha Man came from a bamboo stump.
HEIZMAWNNIA (HEIZMANNIA) PROXIMA n. sp.
(Figs. 33,0; 31,¢ terminalia, claws; 34, pupa, larva)
This species is known only from 2 adult males with associated larval
and pupal skins, It is one of several species (chengi, persimilis n. sp. and
scanloni n. sp. are others) which may previously have been confused under the
name gveenii.
FEMALE. Not known.
34 Conti te oiaees el baeba eiis 54 neat dOT0
MALE. Head. Top of head with pale scales confined to ventrolateral
patches and a median triangular spot between eyes; basal flagellomeres with a
conspicuous spot of pale scales on inner surface; terminal and subterminal
joints greatly elongated; proboscis slender, approximately equal in length to
fore femur; clypeus bare; palpus about one-eighth of the length of the probos-
cis. Thorax, Scutum covered with very broad metallic scales with dark blue
and green reflections; prescutellar bristles absent; scutellum with dark scales
only; anterior pronotal lobes dark scaled except round anterior border; poste-
rior pronotum with pale scales only; no postspiracular bristles; postnotum
with a tuft of pale bristles. Wing. All plume scales very narrow; cell Ro
about 1.8 -2.0 times the length of its stem. Legs. Hind femur pale below,
on anterior surface, nearly to tip; fore tarsus with one claw simple; the other
with a moderately stout tooth near base; both mid tarsal claws simple. Abdo-
men, TergaI-VI dark above with whitish basal lateral triangle prolonged pos-
teriorly along ventrolateral edges. Terminalia. Distimere very much as in
persimilis n. sp.; apex of basimere with a row of 6 setae, represented in the
available material only by their sockets; subapical lobe with a single stout
spine; distal claspette (cl) with 2 pilose lobes; without harpago, very much as
in persimilis n. sp. and propinquan. sp.; proximal claspette (cl) as in
persimilis n. sp. but with all setae in the main tuft approximately equal in
length and with about 5 of them stouter and darker than the remainder; a small
separate lobe; with 2 long setae, present, as in persimilisn. sp. and propin-
quan. sp.; paraprocts with 2 coarse teeth at tip; IXth tergum not well seen,
apparently much as in persimilis n. sp. and propinqua n. sp.; [Xth sternum
completely sclerotized; a stout bristle arising near apex.
PUPA. Cephalothorax. Seta 6-C shorter than 7-C, single; 7-C sin-
gle or bifid. Abdomen. Seta 1-II relatively inconspicuous, with about 1-3
primary and several delicate secondary branches; 5-II about equal in length to
2-II or longer, but much more slender; 9-II-VI minute; 5-IV about equal in
length to the following segment; 5-V somewhat longer; 5-VI about half the
length of the following segment; all of these single; 9-VII and 9-VIII single and
single or bifid respectively. Paddle, About 2.1 - 2,2 times as long as its
greatest breadth; fringe relatively short and scanty; seta 1-P single; male
genitalic sac with apical horns clearly differentiated.
LARVA. Head, Antenna moderately long, curved, spiculate; seta
1-A with 3-4 branches; mouthbrushes without pectinate setae; seta 5-C with
6-7 branches; 6-C single, expanded just beyond base, or with 2 very unequal
branches. Abdomen. Comb a patch of 14-22 spines with greatly hypertro-
phied median denticle; siphon about 3.5 times the length of the saddle (about
4,5 times its breadth at base); pecten of 5-9 slender teeth, all or most of them
with small or very small secondary denticles at base; seta 1-S with 2 slender
branches, arising just beyond distal pecten tooth; saddle with upper portion of
distal edge strongly spiculate; seta 1-X single; 2-X bifid; 3-X single; ventral
brush with 6 setae in the barred area, all single or bifid, and 1 smaller pre-
cratal seta.
TYPE DATA. Holotype male on pin, with terminalia on slide, in U.
S. National Museum; type locality: Doi Inthanon, Chiang Mai, THAILAND.
Mattingly: Heizmannia in Southeast Asia 35
DISTRIBUTION. Specimens examined: THAILAND, Chiang Mai, Doi
Inthanon 2 males with associated larval and pupal skins. Other Distribution.
Two whole larvae from Ban Thuan Lek and Ban Pha Man, respectively, and 9
whole larvae from Ban Wang Mo, THAILAND, may belong to this species or a
closely related form (see below).
TAXONOMIC DISCUSSION. This species is evidently closely related
to persimilis n. sp. and propinqua n. sp., differing from the former in the
ornamentation of the proximal claspette of the male, from the latter in the
male distimere and from both in the shape of the larval comb spines. The
larvae from Ban Thuan Lek etc., noted above, key to the present species but
differ as follows: mouthbrushes dimorphic, either with or without pectinate
setae; antenna longer in the second case (cp. communis, p. 15); seta 1-A sin-
gle to tetrafid; 5-C with 3-11 branches; 6-C trifid on one side of one larva,
the branches in most cases subequal; comb with 12-18 teeth; siphon 2.6 - 3.2
times as long as saddle; pecten with 4-10 teeth; secondary denticles minute or
absent in most cases; seta 1-S bifid or trifid; distal edge of saddle smooth or
almost so.
BIOLOGY. Holotype and paratype bred out from a tree hole. The
other larvae came from a split bamboo, bamboo stump and stump hole.
HEIZMANNIA (HEIZ MANNIA) REIDI MATTINGLY
(Figs. 35, 9;36,°? terminalia, “claws; 37, pupa, larva)
Heizmannia reidi Mattingly 1957, Cul. Mosq. Indomalay. Area 2: 38 (o*, ?*).
Phoniomyia indica Leicester 1908, Stud. Inst. med. Res. F. M. S. 3(3): 253;
Edwards 1922, Indian J. med. Res. 10: 447; Barraud 1929, Ibid.,
17: 265 and 1934, Fauna Brit. India (Dipt.) 5: 303, mec Theobald.
Heizmannia cheni Lien 1968, Trop. Med., Nagasaki,10: 135 (o'*, 9*, P*, L*).
NEW SYNONYMY.
This is a particularly widespread and common species. It is readily
distinguished by the unusually long, slender proboscis and long upper fork
cell. The larvae, also are quite unlike those of any other species, particular-
ly as regards the comb.
FEMALE. Head, Eyes well separated behind; top of head with the
usual basal lateral white patches and patch of white scales between eyes; eye
margins without white scales; basal joint of antennal flagellum with pale scales
and a few dark ones; proboscis about 1,2 - 1.4 times as long as fore femur;
palpus about one-tenth to one-eighth of the length of the proboscis; clypeus
bare. Thorax. Scutal scales with metallic reflection, tinged, in some speci-
mens, mainly with blue, in others mainly with red; no prescutellar bristles;
scutellar scales usually all dark (but see below under Taxonomic Discussion);
anterior pronotal lobes with a large patch of silvery scales occupying most of
the upper surface; a few dark scales behind and at inner corner; posterior
pronotum dark scaled; postspiracular bristles usually absent; a single pale
bristle sometimes present; postnotum with a well developed tuft of dark
36 Contrib. Amer. Ent. Inst., vol. 5, no. 7, 1970
bristles. Wing. Plume scales broad or moderately broad; cell Ro about 2,7 -
3.7 times as long as its stem. Legs. Hind femur pale below on basal five-
sixths or more of anterior surface with dark dorsal stripe reaching to base.
Abdomen. First tergum with laterotergite entirely pale, otherwise dark; re-
maining terga above with basal lateral triangular pale patches, reaching the
distal edge of the tergum along the ventrolateral border only on tergum II,
sterna with dark apical bands except on II. Terminalia. Postgenital plate
rounded at apex; postatrial plates strongly sclerotized.
MALE. Head, Antenna normally as in other species of the genus
(but see under Taxonomic Discussion). Legs. Both fore tarsal claws strong-
ly toothed at base; one with a longer tooth than the other; both mid tarsal claws
simple. Terminalia, Distimere complex, with a well developed, transparent,
lightly pilose 'tail'; apex of basimere with a row of 3-4 short, flattened, stri-
ated setae; subapical lobe with a single stout spine; distal claspette with a well
developed harpago having a finely attenuated appendage, and with a fan-shaped
bunch of short, slightly modified setae bent at the tips; proximal claspette with
a dense patch of moderately long, fine setae and a tuft of longer, somewhat
flattened setae with tips recurved; paraprocts with attenuated tips; [Xth ster-
num strongly sclerotized over almost the whole of its area but with a distinct
median lobe.
PUPA. Cephalothorax. Seta 6-C single, shorter than 7-C which is
Single to tetrafid. Abdomen, Seta 1-II with delicate primary branching, usu-
ally abundant, and some secondary branching; 5-II small and delicate, single
to tetrafid; 9-II-VI minute; 5-IV from 0.6 - 1.7 times the length of the follow-
ing segment; all 3 single; 9- VII single to pentafid; 9- VOI with 3-13 branches.
Paddle. About 1.5 - 3.0 times as long as its greatest breadth, with long
fringe of hair-like spicules and inside this a multiple row of shorter spicules;
seta 1-P single, very occasionally split at tip; male genital sac usually with
well marked horns.
LARVA. Head, Antenna long, slender, curved, usually varying
only slightly with the type of mouthbrushes; spiculate for most of their length;
seta 1-A with 3-8 branches; mouthbrushes either with or without pectinate
setae (see under Taxonomic Discussion); seta 5-C with 5-10 branches; 6-C
with 2 unequal branches, rarely single or trifid. Abdomen. Comb a patch of
about 30-60 scales, uniformly fringed at apex; siphon about 2,.5-4.0 times as
long as its breadth at base, about 2.4 - 3.0 times the length of the saddle;
pecten of 5-15 stout teeth, each with a conspicuous basal denticle, with or
without additional smaller denticles; seta 1-S arising just beyond distal pecten
tooth, with 4-8 branches, somewhat longer than diameter of siphon at point of
attachment; saddle with distal edge almost entirely smooth; seta 1-X various-
ly developed, single to tetrafid; 2-X with 3-7 branches; 3-X single; anal papil-
lae relatively narrow, very variable in length, the upper pair slightly longer
than the lower, the apices pointed; ventral brush with 8 multi-branched setae,
7 of them, at least, with barred sockets, 1-2 of the anterior ones often dis-
placed posteriorly and arising level with, and to one side of, the main barred
area in both Malayan and Thailand specimens,
Mattingly: Heizmannia in Southeast Asia 37
TYPE DATA. 4d. veidi Mattingly, holotype male on pin in British
Museum; type locality: Sukna, N. Bengal, INDIA; allotype female on pin in
British Museum; type locality: Tindharia, N. Bengal, INDIA. H. cheni Lien,
holotype male and allotype female, with associated larval and pupal skins, in
Taiwan Provincial Malaria Research Institute, Taipei, Taiwan; type locality:
Feng-chihu, Chuchi Township, Chiai, TAIWAN.
DISTRIBUTION. Specimens examined: INDIA, North Bengal, Sukna
1 male, Tindharia 1 male, 4 females, Marriambarrie 1 female, Kurseong 1
male, Sureil 2 males. UPPER BURMA, Nam Tamai Valley (27 42' N., 97 54'
E.) 1 female. THAILAND, Chanthaburi, Khao Saidao 8 males, 3 females, 4
whole larvae, 6 larval skins, 9 pupal skins, Ban Thuan Lek 3 males, 3 fe-
males, 3 whole larvae, 4 pupal skins, Chaung Khao 1 whole larva; Nakhon
Nayok, Pha Klory Mai 4 males, 3 females, 1 whole larva, 7 larval skins, 8
pupal skins, Muang 1 female; Lamphun, Kang Arb Nang 1 female; Nakhon Si
Thammarat, Khao Luang Banna 5 females, Khao Keo Banna 3 females; Chiang
Mai, Doi Suthep 7 females, 4 males, 9 whole larvae, 3 larval skins, 4 pupal
skins; Phatthalung, Muang 1 male, 6 females, 6 larval skins, 7 pupal skins;
Tak, Khao Salak Phra 1 male, 1 larval skin, 1 pupal skin; Nakhon Ratchasima,
Korat (Pak Chong) 2 females, 1 male, 3 whole larvae, 1 pupal skin; Phangnga,
Doi Sam Sao 1 male, 3 female, 1 larval skin, 3 pupal skins, Thapwen 6 fe-
males, 1 male, 2 larval skins, 7 pupal skins, Pathum Thani 2 females, 2
males, 2 larval skins, 2 pupal skins; Ranong, Ban Chatri 22 females, 21
males, 42 pupal skins, 10 larval skins. WEST MALAYSIA, Selangor, Ulu
Gombak, 4 males, 16 females, 8 larval skins, 8 pupal skins, Kuala Lumpur
(Pahang Road) 1 male, 1 female. VIETNAM, Long Khanh, Xuan Loc 2 fe-
males, 2 larval skins, 2 pupal skins. TAIWAN, Chuchi 2 males, 2 females
with associated larval and pupal skins. Other Distribution. Probably the spe-
cies recorded by Causey (1937: 414) from northern THAILAND as indica, Ad-
ditional records from TAIWAN include Fanlu, Paiho and Hsiulin (Lien 1968).
TAXONOMIC DISCUSSION. dH. cheni appears to be identical in all
significant respects with the present species. H. indica, confused with it by
previous authors, is quite different (Mattingly 1957 and see p. 25). The distal
claspette and paraprocts recall macdonaldi but this differs in many other re-
spects. A specimen from Kuala Lumpur with male terminalia and female an-
tennae is presumably a gynandromorph. Mouthbrush dimorphism is common
though associated modifications of the antenna are not well marked. A few
larvae with very strongly pectinate mouthbrushes have unusually short anten-
nae. In some Malayan specimens a few scales on the mid lobe of the scutel-
lum appear pale in certain lights. One specimen from Vietnam has a white
spot in this position. The other has the scutellum rubbed.
BIOLOGY. The Malayan larvae were obtained from eggs laid by wild
caught females in the laboratory. Breeding places in Thailand include tree
holes, bamboo internodes, a bamboo pot, a banana tree, ground pools, a rock
hole and acrab hole. The Vietnam larvae came from a tree hole, those from
Taiwan from a bamboo stump. Adults have been taken in a light trap and on
man,
38 Contrib. Amer. Ent. Inst., vol. 5, no. 7, 1970
HEIZMANNIA (HEIZMANNIA) SCANLONI na. Sp.
(Fig. 38, o& terminalia)
This species is known only from a unique male adult, in very poor
condition except for the terminalia which are quite different from those of any
other species. It is one of several species which may previously have been
recorded from Thailand under the name of greeniv.
FEMALE, Not known.
MALE. Head. Top of head denuded; antennae missing; proboscis
broken before tip but apparently long and slender; clypeus bare. Thorax.
Scutum and scutellum entirely denuded; anterior pronotum and posterior pro-
notum denuded except for a few broad white scales which suggest that both may
be extensively pale scaled; some dark scales posteriorly on anterior pronotum;
postnotum with some very pale bristles and a single broad scale. Wing. Great-
ly denuded, remaining plume scales all narrow or very narrow; cell Ro about
2 times as long as its stem or slightly less. Legs. All tarsi and the whole of
both hind legs missing. Abdomen. Missing except for the terminalia which
were remounted ona slide. Terminalia. Distimere as figured; apex of basi-
mere with a dense tuft of flattened, contorted setae with finely attenuated tips,
some pointed, other blade-like, still others racket-shaped; subapical lobe
with 2 stout spines, one slightly smaller than the other; proximal claspette
with dense tuft of long, flattened setae; distal claspette broad, membranous,
pilose with 2 arms, one with a group of short spines at base and a single more
delicate spine near apex (possibly representing the harpago), the other with a
blunt sclerotized appendage somewhat as in persimilis n. sp. (p. 30); apex of
paraproct with 4 small teeth; [Xth tergum with well separated lobes, one with
3, the other with 4 fine hairs; [Xth sternum extensively sclerotized but with
well defined broad median lobe and, apparently, 3 bristles behind apex.
PUPA and LARVA. Not known.
TYPE DATA. Holotype male, on pin, with terminalia on slide, in U.
S. National Museum; type locality: Ban Chang Kien, Chiang Mai, THAILAND.
DISTRIBUTION. Known only from the holotype. |
TAXONOMIC DISCUSSION. Despite the poor condition of the only
available specimen, there can be no doubt, from the terminalia, that this is a
very distinct species. The remarkable flattened setae at the apex of the basi-
mere recall achaetae and, to a less extent, aureochaeta, as does the feeble
development of postnotal hairs. Aedine features are less pronounced than in
achaetae, but certain features of the distal claspette recall subgenus Pavaedes.
It is possible that the early stages, when discovered, may share some of the
features of achaetae and aureochaeta. The female, when found, may well
prove to have some loosely attached broad scales on the postnotum, as in
complex. The only scale remaining in this position in the holotype does not
appear to have arrived by transport.
BIOLOGY. The only known specimen was taken in a light trap ina
garden.
Mattingly: Heizmannia in Southeast Asia 39
HEIZ MANNIA (HEIZMANNIA) SCINTILLANS LUDLOW
(Figs. 1,0; 39,09; 40, ¢? terminalia, o claws; 41, pupa, larva)
Heizmannia scintillans Ludlow 1905, Canad. Ent., 37: 130 (9).
Heizmannia pilosa Brug 1931, Tijdschr. Ent., 74: 245 (o*).
This is a widespread and common species. Females are distinguish-
able from those of other species, except funerea and indica by the prolongation
of the apical dark band on the hind femur ventrally, as well as dorsally, to-
wards the base. Females of indica appear to differ mainly in the predomi-
nantly blue reflection from the scutal scaling and the truncated basal lateral
pale triangles on the abdominal tergites. Females of funerea differ in having
the pale spot on the anterior border of the anterior pronotal lobes almost com-
pletely suppressed. Both differ from scintillans in the male terminalia. The
larvae of scintillans and funerea appear to be indistinguishable and those of
persimilis and propinqua also resemble them closely. The larva of indica is
unknown.
FEMALE. Head, Top of head dark except for the pale ventrolateral
patches and pale spot between eyes; no pale scales round eye margins; basal
joint of antennal flagellum with pale scales and a few small, dark ones; pro-
boscis slender, about 1.1 - 1.2 times as long as fore femur; clypeus bare;
palpus about 0.1 - 0.2 of the length of the proboscis. Thorax, Scutal scales
with metallic- gold reflection tinged with red, blue and green, usually with
blue and green predominating, occasionally red; prescutellar bristles absent;
scutellum entirely dark scaled except in one specimen from Malaya and three
from Vietnam which have a small spot of white scales at the apex of the mid
lobe; anterior pronotum with a variable amount of pale scaling, sometimes
restricted to the anterolateral corner, sometimes covering most of the lobe,
but always with some dark scaling behind; posterior pronotum wholly dark;
postspiracular bristles absent; postnotum with a well developed tuft of bristles.
Wing. Plume scales mostly narrow, broader on Rg and Rg; cell Roy normally
about 2.0 - 2.5 times as long as its stem (see under Taxonomic DiScussion).
Legs. Hind femur with a well developed dark dorsal stripe extending to base
and with a dark ventral stripe variously developed; sometimes failing to reach
base; apical 0.2 - 0.3 of anterior surface entirely dark; first hind tarsal seg-
ment generally pale for some distance below though less so than in the male;
all tarsal claws simple. Abdomen, Tergum I dark; laterotergite white scaled
terga II-VI dark above with triangular basal lateral pale patches prolonged in-
wards but not meeting in mid line; VIL with or without a basal pale bandor spot;
the basal lateral pale spot on II prolonged along the ventrolateral border to the
posterior edge of the segment; the corresponding spot less extensively pro-
longed, if at all, on the other segments; sternum II all pale or with narrow
dark apical band; II-VI or sometimes VII with broad dark apical bands. Ter-
ate ns Postgenital plate rounded at apex; postatrial plates strongly devel-
oped.
MALE. Head, Antenna of normal male type with the 2 terminal seg-
ments greatly elongated. Legs. Second hind tarsomere with a longitudinal
40 Contrib. Amer. Ent. Inst., vol. 5, no. 7, 1970
row of sub-erect scales; third hind tarsal segment with a conspicuous row of
erect scales, becoming sub-erect towards apex of segment; fore tarsus with
one claw simple, the other with a small tooth at base; mid tarsal claws both
simple. Terminalia, Distimere complex; apex of basimere with a row of
conspicuous, flattened, leaf-like setae; subapical lobe with a single spine;
distal claspette recalling that of indica (p. 25) but neither group of setae with
hooked tips; proximal claspette also recalling that of indica but much more
strongly setose and without the fimbriated appendage characteristic of that
Species; paraprocts strongly sclerotized and toothed; [Xth sternum completely
sclerotized.
PUPA. Cephalothorax. Seta 6-C single; 7-C less than 2 times as
long as 6-C, single to trifid. Abdomen. Seta 1-II with expanded base and
fine primary branches which may be numerous but which show little or no
secondary branching; seta 5-II small, delicate, single or bifid; 9-II-VI minute;
5-IV from 0.7 - 1.5 times as long as the following segment, single; 5-V from
1,1 - 1.5 times as long as the following segment, single, occasionally split
at tip; 5-VI from 0.7 - 1.2 times as long as the following segment (0.5 times
in one specimen), also occasionally split at tip; 9- VII single in all cases; 9-
VIII single to tetrafid. Paddle. From about 1.8 - 2,5 times as long as its
greatest breadth; fringe a double row of moderately long hairs with some
shorter ones; a few very small spicules inside the fringe; seta 1-P single, oc-
casionally split at tip; male genital sac frequently bluntly rounded at apex,
without the apical horn characteristic of some other species.
LARVA. Head, Antenna variable, shorter in larvae with pectinate
mouthbrush setae (cp. reidi, p. 35); seta 1-A single or bifid, rarely trifid,
in larvae with pectinate mouthbrushes, with 2-6, usually 3-6 branches in
those with filter feeding mouthbrushes; mouthbrushes dimorphic, with inner
setae pectinate in all but 2 of the Malayan larvae, non-pectinate in all but 2
of the Thailand larvae and in those from Singapore and Sumatra; seta 5-C with
3-16 branches, usually with 10 or more; 6-C normally with 2 unequal branches,
occasionally single, or with one of the branches split, on one side. Abdomen.
Comb with 10-24 mixed scales, with uniformly fringed apex, and spines with
hypertrophied median denticle; siphon about 2-4 times as long as saddle
(about 2 - 4,5 times as long as its breadth at base); pecten with 2-9 large,
simple teeth; secondary denticles minute or absent; seta 1-S single to trifid,
longer than diameter of siphon at point of attachment; saddle with distal edge
at least lightly spiculate; seta 1-X single or bifid; 2-X bifid, rarely trifid;
3-X single; anal papillae large, cylindrical, bluntly rounded at tip; ventral
brush with 6 setae in barred area and 2 precratal setae, all either single or
bifid.
TYPE DATA, d. Sscintillans Ludlow, holotype female, on pin, in U.
S. National Museum; type locality: Camp Stotsenberg, Angeles, Pampanga,
Luzon, PHILIPPINES. 4. pilosa Brug, holotype male, on pin, with termina-
lia on Same mount and wing and associated larval and pupal skins on slides in
British Museum; type locality: Dermajoe, near Benkoelen, Sumatra, INDO-
NESIA,
Mattingly: Heizmannia in Southeast Asia 41
DISTRIBUTION. Specimens examined: WEST MALAYSIA, Selangor,
Ulu Gombak 13 males, 26 females, 20 pupal skins, 20 larval skins, Kuala
Lumpur 1 female, Puchong Road 1 male, 1 female, 2 pupal skins, 2 larval
skins; Perak, Cameron's Highlands 1 female; Pahang, Kuantan 2 males. EAST
MALAYSIA, Savawak, Kuching 3 females, Long Tebanggan 1 female, Gunong
Matang 1 female; Kalimantan, Tarakan 1 female. SINGAPORE 1 male, 1 fe-
male, 1 pupal skin, 1 larval skin, INDONESIA, Sumatva, Dermajoe 2 males,
2 pupal skins, 2 larval skins. PHILIPPINES, Unnamed locality 1 female;
Leyte, Mt. Lob 1 female; Luzon, Calaccad 2 females, Camp Stotsenberg 1
female, Los Banos 1 female, Mt. Isarog 1 male; Mindanao, Dansalan 1 fe-
male, Cagayan 1 female; Palawan, Puerta Princesa 3 females. THAILAND,
Nakhon Nayok, Pha Klory Mai 2 males, 2 females, 6 pupal skins, 2 larvae,
1 larval skin, Sariga Village 1 female, Khao Yai 1 male, 1 female; Nakhon
Ratchasima, Korat 2 males, 11 females, 34 pupal skins, 51 larvae, 23 larval
skins, Nakornsajasima 1 female; Songkla, Boriphat Waterfall 1 female; Ranong,
Unnamed locality 1 female; Trat, Ko ChangIs. 2 females, Lam Ncob 1 male;
Chanthaburi, Chaung Khae 1 larva; Chiang Mai, Ban Patok 1 female; Khon
Kaen, Padong Larn 3 females; Phatthalung, Unnamed locality 1 female; Chum-
phon, Ranong 1 larva; Chon Buri, Bang La Mung 1 female. VIETNAM, Con
Son 27 females. CAMBODIA, Kirirom 2 males. Other Distribution. This is,
perhaps, the species recorded by Borel (1930), as greenii from Xuan Loc,
VIETNAM.
TAXONOMIC DISCUSSION. The discovery of the male from Mt.
Isarog, Luzon, the first from the Philippines, has confirmed the synonymy
previously arrived at on the basis of females only. Records from Vietnam
still require confirmation from males and early stages. Three females from
Con Son have a small spot of white scales at the apex of the mid lobe of the
scutellum but so do 2 females from Thailand so that this may be a variable
character as in some other species. The ratio of cell R, to the length of its
stem is constant within the limits stated except that the Specimen from
Cameron's Highlands has a ratio of about 3 times while the specimen from
Gunong Matang has a ratio of about 1.75 times. The reflections from the scu-
tal scales are usually blue, green and gold but some Malayan and Vietnamese
specimens have them strongly tinged with red. (cp. veidi
BIOLOGY. Adults bite man in forest. Breeding places includes tree
holes, bamboo internodes and a banana axil.
HEIZMANNIA (HEIZMANNIA) TAIWANENSIS LIEN
(Figs. 5, 9; 42,0? terminalia, « claws; 43, pupa, larva)
Heizmannia taiwanensis Lien 1968, Trop. Med., Nagasaki, 10: 137 (o*, 9*,
p*
, L*).
This species is related to greenii, lii Wu and kanhsiensis Tung. Dif-
ferences are noted below under Taxonomic Discussion.
42 Contrib. Amer. Ent. Inst., vol. 5, no. 7, 1970
FEMALE. Head. Top of head with patches of broad white scales at
sides and between eyes, the former extending for a short distance round eye
margins; basal joint of antennal flagellum with pale scales; palpus dark, be-
tween 0.1 - 0. 2 the length of the proboscis; proboscis slender, slightly
shorter than fore femur, dark with a small pale spot below at extreme base.
Thorax. Scutal scales with dull bronze reflection; some pale scales in pre-
scutellar area and on mid lobe of scutellum; lateral lobes of scutellum entire-
ly dark scaled; a small pale spot in front of wing root; postnotum with a group
of pale setulae and 2 broad pale scales; anterior pronotal lobes pale scaled
anteriorly, narrowly dark behind; posterior pronotum covered with broad pale
scales. Wing. Plume scales narrow; cell Ro about 3.0 - 3.5 times as long
as its stem. Legs. Hind femur pale anteriorly on about the basal four-fifths
or more with a dark ventral stripe and a broad dark dorsal stripe both reach-
ing to base. Abdomen. Laterotergite pale scaled; terga II-VI with basal lat-
eral pale triangles prolonged backwards along ventral edge, reaching posteri-
or edge of tergite only on II and III; these patches also prolonged inwards to-
wards mid line, forming a complete pale basal band on VI; tergum VII with a
narrow basal pale band; terga III-V with or without median basal pale spots;
sterna not seen. Terminalia, Without special features.
MALE. Head, Antenna with the 2 distal joints greatly elongated;
verticillary hairs scanty. Legs. Fore tarsal claws subequal; one with astout
tooth arising near base; the other simple; one mid tarsal claw simple; the
other represented by a curved, striated scale-like structure. Terminalia.
Distimere without special features, much as in communis and macdonaldi;
apex of basimere with tuft of dark, flattened, twisted setae as in gveenii and
its allies; subapical lobe with 2 spines; appendage to harpago more or less
falciform; proximal claspette with tuft of very long setae and 2 tufts of shorter,
spiculate setae; paraprocts long, slender, attenuated as in macdonaldi; IXth
tergite with 2 separate lobes, one with 3, the other with 4 setae; IXth sternum
rather elongated with narrow, well defined median lobe.
PUPA. Cephalothorax. Seta 6-C shorter than 7-C, single; 7-C bifid
or trifid. Abdomen. Seta 1-II with 2-4 primary branches, 1 or 2 of them with
secondary branching; 5-II short, slender; 9-II- VI minute; 5-IV and 5-V single,
about as long as the following segment; 5-VI much shorter, single or bifid;
9-VII single or bifid; 9- VIII single to tetrafid. Paddle, Fringe long though
relatively scanty; male genital sac with apical horns relatively broad and
blunt; seta 1-P single.
LARVA. Head, Antenna slender, curved, lightly spiculate, pigment-
ed distally; seta 1-A with 2-3 plumose branches; inner mouthbrush setae slen-
der, simple (occasionally strongly pectinate according to Lien 1968); seta 5-C
with 9-10 branches; 6-C with 2-3 branches, one longer and stouter than the
other(s). Abdomen. Comb a patch of 14-17 uniformly fringed scales; siphon
about 4-5 times as long as saddle with seta 1-S long, bifid or trifid, arising
just beyond the terminal pecten tooth; pecten with 6-10 large teeth, each with
1 or more well developed secondary denticles; saddle with apex finely spicu-
late; seta 1-X bifid or trifid, 2-X bifid or trifid, 3-X single; ventral brush
with 8 setae, all or all but 2 of them in the barred area and all single or bifid;
anal papillae very long, broad, bluntly rounded at apex.
Mattingly: Heizmannia in Southeast Asia 43
TYPE DATA. Holotype male and allotype female, with associated
larval and pupal skins, in Taiwan Provincial Malaria Research Institute,
Taipei; type locality: Kuantzulin, Paiho Township, Tainan, TAIWAN.
DISTRIBUTION. Specimens examined: 1 male and 1 female para-
types, with associated larval and pupal skins, from type locality. Other Dis-
tribution. Said to be widely distributed, but rare, in the whole of TAIWAN
south of latitude 24940'N.
TAXONOMIC DISCUSSION. This species is clearly related to greenii,
from southern India and Ceylon, and to its allies /z7, from China and Korea,
and kanhsiensis from China. H.lii differs in having numerous very broad,
leaf-like setae at the apex of the basimere and both this and greeniz (and, ap-
parently kanhsiensis) differ in the appendage to the harpago. H. greeniihas
a small pale spot below at the base of the proboscis. JH. lii has the proboscis
all dark. The present species also resembles communis and macdonaldi
which share with it, and with the gveenii group, the double spine on the sub-
apical lobe of the basimere. Finally the spiculate setae on the proximal clasp-
ette furnish an interesting link with achaetae and subgenus Mattinglyia.
The curious mid tarsal claw of the male may be an aberration. It is
Shown by both available specimens but these may have come from the same
egg batch. In 177 both mid tarsal claws are simple and nearly straight. In
green each has 2 delicate teeth arising near the base.
BIOLOGY. Breeding places are said to include bamboo stumps and
tree holes below 1000 m. Adults are said to bite man during the daytime in
forest (Lien 1968).
SUBGENUS MATTINGLYIA LIEN
Mattinglyia Lien 1968, Trop. Med., Nagasaki, 10: 128. Type species:
Mattinglyia catesi Lien.
This was erected by Lien (1968) as a distinct genus. However,
thelmae n. Sp. (p. 48) Shows So many annectent characters that subgeneric
status is clearly more appropriate. The principle subgeneric characters are
as follows:
FEMALE. Anterior pronotal lobes well separated; lower mesepime-
ral bristle present (except in catesi); scutellum with broad, silvery scales on
mid lobe; postnotal bristles and scales absent; fore and mid tarsal claws
ames 5 Goat in thelmae); lateral plates of I[Xth tergite fused (except in
thelmae).
MALE. (Unknown in thelmae). Antenna strongly plumose; distimere
relatively simple; subapical lobe of basimere inapparent, the spine or spines
normally associated with it apparently represented by 2 flattened leaf-like
structures with expanded apices; one lobe of proximal claspette with a fan-like
array of flattened blades, the other with recurved setae.
44 Contrib. Amer. Ent. Inst., vol. 5, no. 7, 1970
HEIZMANNIA (MATTINGLYIA) ACHAETAE (LEICESTER)
(Figs. 1,0; 44, 9; 45, 2 terminalia, o claws; 46, pupa, larva)
Dendromyia achaetae Leicester 1908, Cul. Malaya, 257 (9).
Heizmannia stonei Mattingly, Thurman 1959, Univ. Maryland Agr. Expt. Sta.
Bull. A-100: 72 (misidentification).
Resembles catesi but differing in the characters given in the keys.
FEMALE. Head. Top of head with broad, dark scales and a patch
of silvery scales on either side; a patch of broad, silvery scales on vertex;
eyes widely separated behind; tori almost touching; basal joint of antennal
flagellum with small, dark scales; proboscis moderately stout, somewhat
swollen before tip, exceeding the length of the fore femur by a little more than
the length of the labellum; clypeus bare; palpus about one-sixth of the length
of the proboscis or a little less. Thorax. Scutal scales usually with golden
and reddish purple reflections, occasionally appearing mainly green; prescutel-
lar bristles absent; midlobe ofscutellum withbroad,silvery scales; anterior
pronotal lobes smaller than usual with broad silvery scales; posterior prono-
tum largely or wholly covered with broad silvery scales; sternopleuron cover-
ed with silvery scales except for the prealar area and the anteroventral quad-
rant which are bare; mesepimeron with a similar patch covering most of the
upper three-fourths but with the posterior border bare; postspiracular bris-
tles absent; posterior border of sternopleuron with a single long, stout bristle;
postnotum without scales or bristles. Wing. Plume scales on veins Ro and
Re broad; cell Ro always less than 2,5 times as long as its stem, aeually less
than 2times. Legs. Hind femur pale anteriorly and posteriorly on about the
basal three-fifths or less, the dark dorsal stripe very narrow, sometimes in-
complete; fore and mid tarsal claws each with a single small tooth. Abdomen.
Tergum I dark; tergum II with a basal lateral patch of silvery scales, not pro-
longed posteriorly along ventral border; terga IIJ- VII with similar basal later-
al patches but narrower and prolonged inwards towards mid line; V-VII with
continuous silvery basal bands, sometimes incomplete or, rarely, absent on
V and VI; sternum VII dark; the other sterna with silvery basal bands. Ter-
minalia, Cerci broader than usual; postatrial plates strongly sclerotized,
each with a strongly spiculate area at its inner edge; [Xth tergum a single
broad bilobed plate.
MALE. Head, Antenna (Figure 1) with the 2 terminal joints elon-
gated and with short hairs; remainder of flagellum strongly plumose; palpus
as in female. Legs. Fore and mid tarsi with one claw toothed at base, the
other with a smaller tooth at half way. Terminalia. Distimere slender, bent
inwards through approximately 90 degrees, with a prominent flange and stout
Sspiniform appendage; apex of basimere with a dense cluster of flattened, point-
ed, fimbriated setae; subapical lobe not developed, the stout spines which it
bears in other species of the genus here represented by 2 thickened setae with
apices expanded to form transparent leaflets; these leaflets appearing single
in most preparations but seen to be double on rotation; inner tergal edge of
Mattingly: Heizzmannia in Southeast Asia 45
basimere with several closely packed rows of hooked setae; proximal claspette
(bcl) bilobed, one lobe with numerous fine hairs becoming progressively short-
er anteriorly, the other with broad, flattened setae; distal claspette cl) with
3-6 short, slightly flattened setae; harpago absent; paraproct slender, strong-
ly sclerotized distally, expanded at base; Xth sternum membranous, pilose
basally; aedeagus with lateral plates strongly toothed, rather loosely hinged
and easily separated by pressure, their apices overlapping; IXth tergum
smoothly concave, without lobes; [Xth sternum sclerotized at sides and with
a median sclerotized area bearing a group of stiff bristles.
PUPA. Cephalothorax. Seta 6-C long, relatively stout, single; 7-C
shorter, more delicate, single to trifid. Abdomen. Seta 1-II very feebly de-
veloped, single to trifid; 5-II very small, slender, single; 9-II-VI minute;
o-IV single, of the same order of length as the following segments; 5-V-VI
single, very short, markedly smaller than 6-VI; 9-VII with 3-5 and 9-VIII
with 3-9, plumose branches. Paddle, About 1.6 - 1.8 times as long as broad
with multiple fringe of fine hairs and spicules; seta 1-P usually bifid, occa-
sionally single or trifid; male genital sac more or less oblong with short api-
cal horns. :
LARVA. Head. Antenna short, cylindrical, devoid of spicules or
almost so; seta 1-A single, simple; mouthbrush setae all very short, the inner
ones stout and strongly pectinate; seta 5-C with 6-9 branches; 6-C with 2-3
slender, more or less equal, non-plumose branches. Abdomen. Comb of
0-13 small, sharp-pointed spines (cs), delicately fringed at base, in 1-2 ir-
regular rows; siphon about 4 times as long as saddle (about 2.5 times as long
as its breadth at base); pecten of 1-4 very small teeth (i), each with a large
denticle and some smaller denticles at base; seta 1-S bifid, about 1.5 times
the diameter ofthe siphon at point of attachment, arising before half way, just
beyond the distal pecten tooth; saddle with distal edge smooth; seta 1-X long,
stout, plumose, with 1-4 branches; 2-X with 2-4 branches; 3-X single or bi-
fid; anal papillae broad, bluntly rounded, about 4 times as long as the saddle;
ventral brush with 8 setae arising from a barred area; these setae mostly tri-
fid, one or 2 sometimes with 4-5 branches, the 2 shortest often bifid.
TYPE DATA. Holotype female, on pin, in British Museum; type
locality: Jungle patch, 5th mile, Gombak Road, Kuala Lumpur, Selangor,
WEST MALAYSIA,
DISTRIBUTION. Specimens examined: THAILAND, Chiang Mai,
Tad Muang Waterfall 1 male with pupal skin and 2 larval skins, Doi Suthep 1
female; Nakhon Si Thammarat, Khao Kaeo Banna 4 females, Khao Luang Banna
2 females, Songkhla (Boriphat Waterfall) 1 female; Chumphon, Ranong (Kra
Buri) 1 female; Nakhon Ratchasima, Korat (Pak Chong) 2 females; Tvat, Ko
Chang I. 1 female. WEST MALAYSIA, Selangor, Kuala Lumpur (15th and 16th
miles, Gombak Road) 6 males, 16 females, 10 pupal and 10 larval skins, (5th
mile, Gombak Road) 1 female, (Ampang Reserve) 1 female; Pahang, Sabai 1
female; Trengganu, Gunong Tebu 1 female. EAST MALAYSIA, Sabah, near
Kalabakan 1 male. INDONESIA, Celebes, Lambarese 1 female. Other
Distribution. Recorded by Causey (1937) from Bon Pin, THAILAND.
46 Contribs Amer. Ent. Inst., vol. 5, no. -7,<1970
TAXONOMIC DISCUSSION. This species was previously known only
from the adult female. It was placed in the sabethine genus Dendromyia by
Leicester, transferred to Haemagogus by Edwards (1922, 1932) and transfer-
red again, to the present genus, by Mattingly (1957), who noted the resem-
blance to Aedes kanarensis Edwards. The latter was subsequently placed in
Aedes subgenus Diceromyia (Mattingly 1959, 1965). Several characters of
the newly discovered male terminalia confirm the affinity of achaetae with
Diceromyia, especially the concentration of modified setae along the inner
border of the coxite and the large group of flattened setae at the apex (shared
with Haemagogus and some Finlaya). Other characters (structure of proxi-
mal claspette, 'streamlining' of style) recall Group A of Stegomyia with which
Ae. kanarensis also shows affinities. It may not be without significance that
the groups mentioned include all the known, and most of the suspected, vectors
of yellow fever. A fuller understanding of relationships must, however, a-
wait the discovery of the males and early stages of other little known annec-
tent species (Ae. discrepans Edwards, Ae. tripunctatus Theobald, Ae. platy-
lepidus Knight & Hull). The aedine features of the male terminalia are not
sufficiently pronounced to justify the removal of achaetae from Heizmannia
and the characters of the early stages are fully consistent with its retention
in this genus, particularly having regard to the annectent character of thelmae
n. Sp.
The male and early stages described by Thurman (1959) under stonez
belong, in fact, to the present species.
- BIOLOGY. The Thailand larvae came from a tree hole and, in one
case, from a fallen coconut leaf. Those from Malaya were raised from eggs
laid in the laboratory. Adults have been taken on man in forest.
HEIZMANNIA (MATTINGLYIA) CATESI (LIEN)
(Figs. 47, 29; 48, 2 terminalia; 49, pupa, larva)
Mattinglyia catesi Lien 1968, Trop. Med., Nagasaki, 10: 130 («*, 9*, P*, L*).
Resembles achaetae, differing in the characters given in the keys.
The third member of the subgenus, thelmae n. sp., differs from both in var-
ious characters annectent to the previous subgenus.
FEMALE. Head. Top of head dark with large silvery lateral patches
and a silvery median spot extending forward between eyes; eyes well separat-
ed; tori almost contiguous; basal joint of antennal flagellum dark scaled; pro-
boscis long, slender, dark, about 1, 2 times the length of the fore femur;
palpus dark, about one-eighth of the length of the proboscis. Thorax. Scutal
scales broad and flat with bright green, gold and reddish reflections; scutellum
with a silvery spot on mid lobe, lateral lobes dark or with some Silvery scales;
postnotum bare; anterior pronotal lobes covered with silvery white scales;
posterior pronotum with a small spot of silvery scales. Wing. Plume scales
narrow; cell Ry about 1.5 times as long as its stem. Legs, Hind femur en-
tirely dark on about the distal half of both anterior and posterior surface, the
Mattingly: Heizmannia in Southeast Asia 47
remainder pale except for a narrow dark dorsal stripe on most of the basal
half; both fore and both mid tarsal claws toothed. Abdomen. Laterotergite ©
with silvery white scales; terga II-VI with silvery white lateral patches at
most slightly prolonged posteriorly along the ventral edge, prolonged well in
towards mid line; one specimen with a continuous silvery basal band on V;
tergum VII with a silvery basal band; sterna with narrow basal silvery bands.
Terminalia, Postgenital plate not deeply indented; [Xth tergum with 1 bristle
on each lobe in the specimen examined.
MALE. Head, Antenna with the 2 terminal joints together about
equal in length to the remainder; the other 11 joints strongly plumose. Legs.
Fore and mid tarsal claws strongly toothed (fide Lien), the former much the
larger. Terminalia. Much as in achaetae but with the 2 subapical leaflets
differently shaped; flattened blades on proximal claspette only 7 in number,
the fine hairs fewer, all of them slender and more or less equal in length;
distal claspette with 1-2 slender setae; setae on inner face of basimere very
strongly hooked, arising from a prominent lobe.
PUPA. Much as in achaetae except that setae 5-IV and 5-V are lon-
ger than the following segments while 5-VI is about two-thirds as long as seg-
ment VII. Paddle. Spiculate, distinctly broader than in achaetae, about 1.5
times as long as its greatest breadth. |
LARVA. Head, Antenna short, straight, smooth; seta 1-A single;
inner hairs of mouthbrushes slender, simple; seta 5-C with 10-11 branches;
6-C bifid. Abdomen. Comb with 15-20 sharply pointed teeth (cs), fringed at
base; pecten of 4-5 teeth (pi), each denticulate on both sides; seta 1-S trifid
or tetrafid; seta 1-X bifid; 2-X tetrafid; 3-X single; anal papillae about 5
times as long as saddle, bluntly rounded at tips; ventral brush with 8 bifid or
trifid setae.
TYPE DATA. Holotype male and allotype female, with associated
larval and pupal skins, in Taiwan Provincial Malaria Research Institute,
Taipei; type locality: Yuehmei, Sanlin, Kaohsiung, TAIWAN.
DISTRIBUTION. Specimens examined: TAIWAN, holotype male;
Pingtung, Taiwu, Mashihshih 3 paratype females. Other Distribution. Re-
corded by Lien (1968) from the type locality and Mashihshih only.
TAXONOMIC DISCUSSION. Although closely related to achaetae
this species is clearly distinct on the characters of the male terminalia and
pupa. In one specimen seen by me the median silver spot on the occiput is
completely fused with one of the lateral silver spots. There is thus a nearly
complete silvery band on the occiput recalling the condition seen in thelmae
n. sp. The complete, or almost complete, basal silver bands on the abdomin-
al tergites also recall that species which differs, however, in important char-
acters of the female terminalia and claws as well as the all dark anterior pro-
notal lobes.
BIOLOGY. The only known breeding places are bamboo stumps. Re-
corded as biting man viciously in the daytime.
48 Contrib. Amer. Ent. Inst., vol. 5, no. 7, 1970
HEIZMANNIA (MATTINGLYIA) THELMAE n. sp.
(Figs. 50, 9; 38, 2 terminalia, claws;51, pupa, larva)
Differs from the other members of the subgenus in the all dark ante-
rior pronotal lobes and from achaetae in the markings of the occiput and ab-
dominal terga. It seems possible, however, that, as noted above, the present
Species and catesi may show overlapping variation in these characters.
FEMALE. Head. Top of head entirely covered anteriorly with broad
flat, silvery white scales, broad dark scales behind; some small, dark up-
right scales on occiput; eyes close together except for a short distance behind;
antenna with small pale scales on basal joint of flagellum; proboscis short and
stout, somewhat swollen towards tip, about 0.8 times as long as fore femur;
palpus nearly a quarter of the length of the proboscis. Thovax. Scutalscales
broad with bright, mainly bluish, reflection and silvery highlights; prescutel-
lar bristles absent; scutellum with a large spot of silvery scales at apex of
mid lobe; anterior pronotal lobes and posterior pronotum entirely dark scaled;
postspiracular bristles absent; postnotum bare. Wing. Plume scales on R
and Re relatively broad especially towards tip; anterior fork cell very long,
about 3.3 times as long as its stem. Legs. Hind femur pale on about the bas-
al two-thirds with very narrow dark dorsal stripe barely reaching to base;
fore and mid tarsal claws simple. Abdomen. Tergum I dark; remaining terga
with broad silvery basal bands, widening a little at sides on posterior seg-
ments; sterna not seen except VII which is dark. Terminalia. Cerci narrow-
er than usual in the genus. Postatrial plates only lightly sclerotized. IXth
tergum with 2 separate plates as in all other Heizmannia except achaetae and
catesi, Spermatheca triple as usual.
MALE. Not known.
PUPA. Cephalothorax. Seta 6-C stouter, but shorter, than 7-C.
Abdomen. Seta 1-II moderately stout, trifid; 5-II slender, single; 9-II-VI
minute; 5-IV-VI single, shorter than the following segment; 9-VII small, sin-
gle; 9- VIII bifid on one side, trifid on the other.
LARVA. Head, Antenna short, smooth, cylindrical; seta 1-A deli-
cate, bifid; mouthbrush setae apparently non-pectinate; seta 5-C with 9-10
branches; 6-C single on one side, with 2 unequal branches on the other. Abdo-
men. Comb of 8 spines with stout median, and delicate lateral, denticles;
siphcn about 3 times as long as the saddle (3 times as long as its breadth at
base); pecten of 2 very small teeth, simple or with minute denticles at base;
seta 1-S missing; saddle edge smooth; seta 1-X long, single; 2-X bifid; 3-X
single; all these simple; anal papillae short, blunt, cylindrical; ventral brush
with 8 setae in barred area, all single or bifid.
TYPE DATA. Holotype female, on pin, with terminalia, claws and
larval and pupal skins on slides, in the U. 8. National Museum; type locality:
Ban Chatri, Ranong, THAILAND. The species is named after Miss Thelma
Ford in gratitude for her beautiful illustrations to this paper.
DISTRIBUTION. Known only from the holotype.
TAXONOMIC DISCUSSION. This species shares with the others in
subgenus Mattinglyia the absence of scales or bristles from the postnotum.
Mattingly: Heizmannia in Southeast Asia 49
It shares with subgenus Hezzmannia s. str. the simple fore and mid tarsal
claws of the female and the separate lateral plates of the female [Xth tergum.
It is thus annectent between the two subgenera though some striking resem-
blances in ornamentation, particularly to catesi, suggest that it is closer to
Mattinglyia.
BIOLOGY. The only known specimen was bred from a bamboo.
ACKNOWLEDGEMENTS
The main sources of material for this paper have been collections
made by the staff of the Medical Research Institute, Kuala Lumpur, in Malaya,
and the SEATO Medical Research Laboratory, Bangkok, in Thailand, together
with earlier collections in the U. S. National Museum and the British Museum
(Natural History). For the Malayan material, and for data relating to it, Iam
greatly obliged to Dr. W. W. Macdonald of the Liverpool School of Tropical
Medicine. Dr. J. C. Lien kindly supplied paratypes and one holotype of his
new species and Dr. J. M. Klein of the Office de la Recherche Scientifique et
Technique Outre-Mer sent material from Cambodia. Dr. Botha de Meillon
and Dr. Alan Stone kindly made available to me the material from Thailand,
now in the Southeast Asia Mosquito Project and the U. S. National Museum.
I am also indebted to Dr. Stone and to Dr. de Meillon and his colleagues of
the Southeast Asia Mosquito Project for much hospitality and assistance in
Washington. The illustrations prepared by Miss T. L. Ford, Mr. Vichai
Malikul and artists of the U. S. Army 406th General Medical Laboratory,
Tokyo, speak for themselves.
LITERATURE CITED
BARRAUD, P. J.
1929. A revision of the culicine mosquitoes of India. Part XXVI.
The genera Heizmannia, Haemagogus, Topomyia and Mega-
rhinus. Indian J. med. Res. 17: 259-280.
1934. Family Culicidae. Tribes Megarhinini and Culicini. The
Fauna of British India, including Ceylon and Burma. Diptera,
Vol. V;, 463 pp. London.
BOREL, E.
19
30. Les Moustiques de la Cochinchine et du Sud-Annam. 423 pp.
Paris.
BRAM, R. A.
1967. Contributions to the mosquito fauna of Southeast Asia. - II.
The genus Culex in Thailand (Diptera: Culicidae). Contr.
Amer. ent. Inst. 2(1): 1-296.
50 Contrib. Amer. Ent. Inst., vol. 5, no. 7, 1970
BRUG» iby: 4
1931. New Culicidae from Sumatra. Tijdschr. Ent., 74: 245-250.
BRUG, 8S. L. and J. BONNE-WEPSTER
1947. The geographical distribution of the mosquitoes of the Malay
Archipelago. Chron. nat. 103: 179-197.
CAUSEY, O. R.
1937. Some anopheline and culicine mosquitoes of Siam with remarks
on malaria control in Bangkok. Amer. J. Hyg. 25: 400-420.
EDWARDS, F. W.
1922. A synopsis of adult Oriental culicine (including megarhinine
and sabethine) mosquitoes. Part II. Indian J. med. Res. 10:
430-475.
1932. GeneralInsectorum, Diptera. Family Culicidae. Fascicle
194, 258 pp. Brussels.
EDWARDS, F. W. and D. H. C. GIVEN
1928. The early stages of some Singapore mosquitoes. Bull. ent.
Res. 18: 337-357.
JOLLY, C. G.
1933. Report on the mosquito survey of Rangoon. Rangoon: Govern-
ment Printer.
LEICESTER, G. F.
1908. The Culicidae of Malaya. Stud. Inst. med. Res. F.M.58. 3:
18-261.
LIEN, J. C.
1968. New species of mosquitoes from Taiwan (Diptera: Culicidae).
Part IV. Mattinglyia catesi, n. gen., n. Sp. and four new
ehee\s ep genus Heizmannia Ludlow. Trop. Med., Nagasaki,
O: £27-153.
LUDLOW, C. S.
1905. Mosquito Notes, No. 3 (continued). Canad. Ent., 37: 129-
135.
MACDONALD, W. W. and R. TRAUB
1960. Malaysian parasites. XXXVII. An introduction to the ecology
of the mosquitoes of the lowland dipterocarp forest of Selangor,
Malaya. Stud. Inst. F.M.S., 29: 79-109.
Mattingly: Heizmannia in Southeast Asia ol
MARKS, E. N.
1958. Notes on Opifex fuscus Hutton (Diptera: Culicidae) and the
scope for further research on it. N. Z. Ent. 2: 20-25.
MARKS, E. N. and E, P. HODGKIN
1958. A new species of Aedes (Finlaya) from northern Australia
(Diptera, Culicidae). Proc. Linn. Soc. N.S.W. 83: 33-39.
MATTINGLY, P. F.
1957. Cul. Mosq. Indomalayan Area. PartII. Heizmannia Ludlow.
57 pp. London.
1959. Cul. Mosq. Indomalayan Area. PartIV. Genus Aedes
Meigen, Subgenera Skusea Theobald, Diceromyia Theobald,
Geoskusea Edwards and Christophersiomyia Barraud. 61 pp.
London.
1965. Cul. Mosq. Indomalayan Area. Part VI. Genus Aedes
Meigen, Subgenus Stegomyia Theobald (Groups A, B and D).
67 pp. London.
McGREGOR, D. D.
1963. Mouthbrush dimorphism in larvae of Opifex fuscus Hutton
(Diptera, Culicidae). Bull. ent. Res. 54: 325-327.
THEOBALD, F. V.
1905. A catalogue of the Culicidae in the Hungarian National Museum.
Ann. hist.-nat. Mus. hung. 3:61-119.
1910. Second report on the collection of Culicidae in the Indian
Museum, Calcutta, with descriptions of new genera and spe-
cies. Rec. Indian Mus. 4: 1-33.
THURMAN, E. B.
1959. A contribution to a revision of the Culicidae of Northern
Thailand. Univ. Maryland Agr. Expt. Sta. Bull. A-100,
182 pp.
Fig.1
4
{aX
i
y;
N
WN
H. (Mattinglyia) achaetae
H.(Heizmannia) scintillans
4, % |
(A/1sAw7a 44
Japan
H.(Heizmannia ) aureochaeta
spermathecal
duct
Fig..3
IXth sternum
SASS
SOS
9
Sri EN hs
EDULE Reed
Cnt:
RLY].
“as
One
Snes
y/
postgenital
| claspette
distal claspette
(dcl)
proxima
fore claw o7
IXth tergum
Ee 0.05mm———_—
mid claw of
PS OOS mm
ical lobe\
ine
and sp
hind claw o&%
paraprocts
subap
IXth tergum
T ° f (ee) SD om)
H (Heizmannia) aureochaeta
H. (Heizmannia) aureochaeta
get
H.(Heizmannia) chengi Lien
oa
-———- 1-O____
H.(Heizmannia) taiwanensis i Si
Fig.6
re apical setae
subapical spines
and lobe
(pel)
basimere
5
=. -
paraprocts
|
heask’ oy J) fa
ae Nai Wa
0.05mm
cercus
postgenital
wats is
plate
proctiger
spermathecal
0.05mm
uct
postatrial
plate
IXth sternum
aedeagus
+—— 0.1mm ————+4
0.1mm
H. (Heizmannia) chengi
fore claw o7
Fea OC. 06am
mid claw o7
— Fe Secs
H. (Heizmannia) chengi
H.(Heizmannia } communis
TANS
it
SS ah
aps
/
hehe aa AG a
Nai AMS
THUS Mens
filet
cercus
mid claw of
0.1mm
ae. i? SN
ak tgenital
Ke postgenita
distimere plate
spermathecal
uct
: postatrial
apical setae plate
subapical lobe
and spines
0.1mm
spermathecae
basimere
IXth tergum
paraprocts
beak ro
IXth sternum
+— 0.1 mm—‘¥+
OAS H. (Heizmannia) communis
ROY
x
7
Ney
NY
SNS
SAN
}
NY
SNQIAS::
Sy
X
2
H. (Heizmannia) communis
H. (Heizmannia) complex
Fig.12 |
+——_0.1 mm —"1
\ 4 IXth sternum \ aif ates \ NG
A postgenital er iN a DOR
> plate PAN AUARIN NBEN LRN
> MUL OM
HN ie
cercus
~~ 3 Le
WS SSS A
SS
\
ANN NN
a
=
ss
SS
AS
YY
\
\
SELES
LLL ELLIE
postatrial
plate
distimere
insula
\0.05mm
subapical
lobe and
aedeagus paraprocts
( pcl)
IXth tergum : fe | 0.05 mm
basimere
H. (Heizmannia) comp/ex sokeihuas
Sw WN
H. (Heizmannia) complex
UB ——— ts)
H.(Heizmannia ) covelli
fore claw of ne
> 0: 05mm...
distimere
apical
; setae
{ | /
subapical
lobe and
spine
mid claw of
postgenital
plate
GaViiny,
\ uN 1) ) if
(\ ‘ Ni Nee Hy }) Ay
i LS | Ass f sia
Se .
ie NI CH BY iH: ‘i i \
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ede
\
h
ai
a
}
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aN
i
x
IXth sternum Xs
-—— 0. mm—. 2 NN
J) aw AA)
ar ae
itt My spermatheca uct
postatrial
plate
¢
IXth sternum
0.1mm
T. I =],
H. (Heizmannia) covel/i
MP
H.
(Heizmannia) funerea
pote eon ae
H.(Heizmannia ) demeiffoni
= |
eae f
et: |
. D o Ot i
oO ero
omnes o a)
= =
A
Be :
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o=—
ratate's Sas
ss
a
H. (Heizmannia) demei/loni
REVS
SS
SAX
LOSS
S
SS
Ciao
IXth tergum
duct
-——_ 0.05mm——_—-
spermathecal
mid claw 9
ng
:
isp
a,
H. (Heizmannia ) funerea
ge
<<
cee
Sa areca
q
~~
i Ge SS EeM\ Lo spermathecal
\ Bar: GMM oD Pe irae
\ BS IXth sternum ae Ale duct
> y F
postatrial
a
\\ RN
Rieti
SUV
"Kcz
ig
We
y midclaw o&
distimere
~ Poa ores Use
proctiger
subapical
spines and
lobe
0.1mm
Uehaifbhaalhadl
H.( Heizmannia) indica E
)
subapical lobe
and spine
terminalia
(lateral aspect
Fig. 22
oe
SSs>
eo.
basimere
terminalia
(dorsal aspect)
ostatrial
late
2 ee aa
© ® Z
a
Bo
oa
rom
ou
S5
o-O
ee
os
»: re}
=
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o
a
”
paraprocts
IXth tergum
mid claw of
IXth sternum
<i es
proctiger
fore claw o7
H. (Heizmannia) indica
x
H.(Heizmannia ) macdonald
rig. 24%
distimere
mid claw of
apical setae pa Osman
subapical lobe
and spines
fore claw o7
spermathecal
ye duct
-——_——_ 0.1 mm ———“
para procts y,
GZ
postatrial
plate
aedeagus
0.05mm
IXth tergum
g
spermathecae
T. | > =
proctiger
H. (Heizmannia) macdonaldi H. (Heizmannia) mattinglyi
te SS CO } 1
PT
H. (Heizmannia) macdonaldi
Wis Haginana
H.( Heizmannia) mattinglyi
H.(Heizmannia) persimilis
%
tas
divi
Sy
b (ao Ny
Oe
ey
distimere
subapical lobe
and spine
IXth sternum
+-———__ 0.1mm ———_1
0.1mm
IXth tergum
basimere
0.1mm
(pel)
paraprocts
postatrial
postgenital plate
plate
x Nt x SS
eb Sa
: SST?
mid claw o7 SA if
Dh SS
fore claw o7%
H. (Heizmannia) persimilis === —a-e=sR=
MP
H. (Heizmannia) persimilis Se
H.(Heizmannia ) propinqua
setae
subapical
lobe and
spine
(pcl)
pesaeees +—— 0.1 mm —+1
AOS
GFE
fore claw of | IXth sternum
distimere
H. (Heizmannia) proxima
apical setae
subapical lobe
and spine
proctiger
IXth sternum
H. (Heizmannia) propinqua
fore claw o%
M
a]
PT
H. (Heizmannia) propinqua Seeman teeth:
H.(Heizmannia ) proxima
H. (Heizmannia) proxima
RE SR, rae oa ae
fzmannia) reidi
H.(He
er AEE asta Chey,
distimere
mid claw o7
K— 0.05 mm ——_J
Q
apical setae
subapical lobe
and spine
(pcl)
fore claw o7
basimere
0.1mm
IXth tergum
IXth sternum
cercus
H. (Heizmannia) reidi
Pe? Hp) th
HM
fi
a. Sere
H. (Heizmannia) reid/
apical
setae
subapical spines
and lobe
IXth tergum
2
cercus ae
IXth sternum ; i f
postgenital Anes
plate Hy
paraproct
7” cedeagus
H. (Heizmannia) scanloni H.
Fig.38
————--—- 0.05mm—_—_—__—1
0.1mm
SS
Wee Ne
HEA) 1 N
eat i
M,
yes
vase
i")
ry lie
4B)
IXth tergum
(Mattinglyia) thelmae
es
‘TS
7) SS
CT
Dinenscscccomnelt
H.(Heizmannia) scintillans
Fig.40
mid claw o7
0.1mm
cercus
postgenital
IXth sternum
plate
ys
yy
VRS ee
NUN Ih
A
: Sc saminda
O.Iimm
ey
ete)
spermathecal
postatrial
plate duct
ee eetze
IXth tergum
subapical
lobe and
spine
|
au Se on <5
: was é x. of RS
0.1mm
basimere
aedeagus
H. (Heizmannia) scintillans
H. (Heizmannia) scinti/lans
Ue a —— la | ———)
4
1
c
= ae
~---
PT
fore claw o”
postgenital
plate
paraprocts
cercus
spermathecal
duct
postatrial
plate
proctiger
apical setae-- el
‘ IXth tergum
distimere
subapical lobe
and spines
0.1mm
IX th tergum +——0.1 mm—+
IXth sternum
H. (Heizmannia) taiwanensis
/wanens/s
ae
H. (Heizmannia) ta
H.(Mattinglyia) achaetae
t+— 0.05 mm—H
Imm ——
IXth sternum.
re claw o%
aacag *
fo
mid claw o7
plate
postgenital
postatrial
cercus
apica
subapical
leaflet
setae
imere
isti
d
aedeagus
iger
proct
IXth tergum
spermathecae
H.(Mattinglyia) achaetae
Vill
‘
H.(Mattinglyia) achaetae
ST SR
Kt —_—_#_———-_ 0.1 mm ——H41
H.(Mattinglyia) cates
eR Sr ad
cercus
postgenital
plate
SEES
Hin
Nd
NW fit WW:
WiAg
IXth sternum
sie sta
: uct
postatrial
plate
distimere 0.1mm
subapical
leaflet
apical setae
IXth tergum
basimere
IXth tergum
a er — st
H. (Mattinglyia) catesi
H.(Mattinglyia) thelmae
H. (Mattinglyia) thelmae
Mattingly: Heizmannia in Southeast Asia
INDEX
103
Valid names are set in roman type, Synonyms are italicized. The
italicized pages are those which begin the primary treatment of the taxon.
Numbers in parenthesis refer to the figures illustrating the species in ques-
tion.
achaetae
Aedes
anopheloides
aureochaeta
bambusa
Bolbodeomyia
catesi
chengi
chent
Christophersiomyia
communis
complex
covelli
demeilloni
Dendromyia
Diceromyia
discrepans
Finlaya
funerea
ereenil
Haemagogus
Heizmannia
himalayensis
indica
kanarensis
kanhsiensis
lii
macdonaldi
mattinglyi
Mattinglyia
metallica
nivirostris
Opifex
1, 2, 4, 6, 7, 8, 9, 11, 12, 15, 19, 38, 43, 44, 46,
47, 48 w ‘44, 45, 46).
2, 46.
15.
i. Suns, 98, Ge Law Tags bs, 28 (2,, 3, 4),
o
9. 11,1,
ie
35, 37.
nae
(8,
2, 38 (11, 12, 13).
3 (14, 15, 16).
9,°10, 21, 23 (17, 18).
46.
3 3 3
Cy 6,
, 208 31, 33, 38, 41, 42, 43.
, 4, 46.
a ay A,
21, 22, 23.
1. Dy 7, 9; 10, 23, 24,25; 26, 35, 37, 39,
22).
2, 6, 7, 9, 43, 44, 46, 48, 49 (47, 48, 49).
anitol ‘Ky 17, 26, 30 (5, 6, ge
Ba Vy oe Pe bes Ay EDy 2p 223) Zot.
9,40, 23, 24, 25, 26226. 29 (16.
6,28, 13.0 Th. Pgh 2h, (es, Boe
30, 32, 305 38, 39, 41, Ad, 46, 48, 49.
3 8, 10, 11, 15, 20, 21, 26, 27, 28, 35, 42,
10).
2, : 4, 5, T, 8, Ov d1lad, £3, 14, .'27,. Bae
28, 29,
19, 20).
26, 28,
40 (21,
‘hee RE 6, & 10, 14. 15,17, Be, 28: 87, 42, 43
5
5. 7s 9; 10, 21, 29 (24,, 26).
, 2, 3, 4, 6, 43, 44, 46, 48, 49.
1
tS, 20s
1, 26, 28.
2. }
104
Orthopodomyia
Paraedes
persimilis
Phoniomyia
pilosa
platylepidus
pr opinqua
proxima
reidi
scanloni
scintillans
Stegomyia
stonet
taiwanensis
thelmae
Tripteroides
tripunctatus
Wyeomyia
Contrib.
Amer. Ent. Inst., vol. 5, no. 7, 1970
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9
8
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1
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TAGs 19; 144) ae:
£O ae, G1) 8271833084, 9S,--38, 39 (27,
10,245 30) 81, 82) 34) 35, 39°30, 31,932),
10, 24, 30, 31, 32, 33 (31, 33, 34).
Ds th Oa Os EOSHZS,: 26, 95, ST, 40, 41 405;
0, 30, 33, 38 (38).
a, OY 10, 44,) 21,22, 23, 24, 25, 2ay498;
39, 40, 41).
9, 10, 41 (5, 42, 43).
8, 10, 43, 46, 47, 48 (38, 50, 51).
Publication dates for Volume 4 of
Contributions of American Entomological Institute
Volume 4, no.
1 Nov. 8, 1968
no. 2 March 13, 1969
no. 3 July 12, 1969
no. 4 May 16, 1969
no. 5 May 5, 1970
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7 } fs) . : is : my: = i oe a % - _ -, zie i ‘4 ‘ i AS j “J aN : 2. ae *y ae Lan tt
net Fibs, ake Sur tied SUA" hs Us Grane a ton Ma oT a ye Ce Ok
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(Continued from inside front cover)
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