No. 153

OCCASIONAL PAPERS
OF THE
CALIFORNIA ACADEMY OF SCIENCES

February 12, 2003

EMBIA
Contributions to the Biosystematics
of the
Insect Order Embiidina

Part 4
Andesembiidae, a New Andean
Family of Embiidina

By
Edward S. Ross

Published by the California Academy of Sciences
San Francisco, California

mé

EMBIA
Contributions to the Biosystematics of the
Insect Order Embiidina

Edward S. Ross collecting the type culture of Andesembia incompta, n. sp., on
an epiphyte-covered trunk of the tree fern Blechnum buchtiensis. Evident is the
great difficulty of finding small embiid colonies in such a densely vegetated
habitat. It should be noted that there are no host plant associations in Embiidina.
Additional specimens of the species were collected in moss on fence posts.
Colombia: 25 km W of Macoa, 2030 m elev., Prov. Putamayo.

OCCASIONAL PAPERS
OF THE
CALIFORNIA ACADEMY OF SCIENCES

No. 153 February 12, 2003

EMBIA
Contributions to the Biosystematics
of the
Insect Order Embiidina

Part 4
Andesembiidae, a New Andean
Family of Embiidina

By
Edward S. Ross

Published by the California Academy of Sciences
San Francisco, California

SCIENTIFIC PUBLICATIONS

Alan E. Leviton, Editor
Katie Martin, Managing editor

© 2003 California Academy of Sciences, Golden Gate Park, San Francisco, California 94118
All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any
means, electronic or mechanical, including photocopying, recording, or any information storage or re-

trieval system, without permission in writing from the publisher.

ISSN 0068-5461

Table of Contents
Part 4. Andesembiidae, a New Andean Family of Embiidina

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Part 4

Andesembiidae, a New Andean
Family of Embiidina

Abstract

Known only from higher altitudes of the Andes,
Andesembiidae comprises the new genus
Andesembia with six new species from Ecuador and
Colombia and the new genus Bryonembia with one
new species from a cloud forest habitat in central
Peru.

Acknowledgments

I wish to thank Dr. Evert I. Schlinger for assis-
tance and companionship during my 1954—55 field-
work in the Andes, as well as David Cavagnaro for
collecting cultures in Ecuador during 1964.

Introduction

Explanation of terminalia and wing symbols:
9 = ninth abdominal tergite; 10 L = left hemitergite
of abdominal somite 10, 10 LP = its process; 10 R=
right hemitergite of 10, 10 RP = its process; MS =
medial sclerite of somite 10; MF = medial flap
(which may include process of 10 R); EP = epiproct
(somite 11); H = hypandrium (ninth sternite), HP =
its process; GO = gonopophysis; LPPT and RPPT =
left and right paraprocts; LC, LC, = segments of
left cercus. Wing veins: C = costa; RBS = radial
blood sinus (RA); RP = posterior radius; MA and
MP = branches of media; Cu = cubitus.

Institutional symbols: BMNH -— British Mu-
seum of Natural History, London; CAS — California
Academy of Sciences, San Francisco; IHNB —
Instituto de Historia Naturales, Bogota, MUSM —
Museo de Historia Natural, Lima, Peru; NMQ —
Museo de Ciencias Naturales, Quito; USNM —
United States National Museum, Washington.

Paratypes: This designation is limited to
topotypic adult males in keeping with my opinion
(Ross, 1956, Systematic Zool. 5:191—92) that
paratypes should as near as possible represent the
characters of the holotype and not the general na-
ture of the species, a never-to-be completed objec-
tive. Topotypic females are designated parallotypes.

In most cases my type series are from a single cul-
ture. Specimens from more distant localities are
treated as “other specimens examined.”

Andesembiidae Ross
new family

Type genus.—Andesembia Ross, new genus, by
present designation.

Name basis.—Reference to occurrence in Andes.

Diagnosis.—Males: Small to moderately large
(body length 6.5—12 mm), slender; uniformly brown,
including antennae and cerci, head darker than body;
always alate. Wings usually large relative to body
proportions. Head usually disproportionately large.
Eyes small. Basal segment of antennae narrow but
flared distad; other segments slender, elongate; 15
to 20 segmented. Mandibles often large, irregular in
form, always apically dentate. Submentum moder-
ately to strongly sclerotized, margins well defined,
sides apically divergent. Wings large; RBS parallel
to C, terminated before wing apex; RP relatively well
sclerotized; MA sclerotized at base only, continued
unforked to wing margin as a line of setae; cross-
veins absent behind RP. Legs very slender; hind
basitarsus very long, slender, without a medial pa-
pilla. Terminalia relatively small, weakly sclerotized.
Tenth tergite cleft to base; in some species the cleft
is broadly membranous, in others the medio-basal
area (MS) is very broad, fused to a narrow 10 R.
Left hemitergite (10 L) often small, margins weak;
its process (10 LP) always simple but at times long
and sinuous; right process (10 RP) short. Medial flap
(MF) a simple arm connected to base of epiproct
sclerite (EP), which is prominent, curved or angled
beneath medial flap; its forward extremity variously
spiculated from coarse and sparse to minute and
dense. Left paraproct (LPPT) a simple, broad scler-
ite, at times micro-echinulate caudo-mesally, never
bearing a process. Right paraproct (RPPT) occasion-
ally as large as left paraproct, however, at times rep-
resented only as sclerotic fragments. Hypandrium

2 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 153

process (HP) a simple, rounded or truncate, distally-
membranous lobe. Basal segment of left cercus (LC, )
gradually, inwardly expanded distad but never dis-
tinctly lobed, but strongly echinulated; distal seg-
ment either articulated or fused to basal segment.
Basal segment of right cercus not strongly sclero-
tized, inner-basal portion weak, depressed by con-
tact with the large right paraproct.

Females.—Uniformly brownish. Hind basitarsus
without a medial papilla; large setae almost entirely
confined to outer side, those of basal half angled
distad, those of apical half directed toward plantar
side.

Discussion.—This category is proposed for an
interesting group of species occurring at high alti-
tudes in the Andes. Were it not for the consistently
non-dentate mandibles characterizing all genera of
Anisembiidae, one might be tempted to place
andesembiid genera in that family. However, the
Andean genera are too distinct to place in any known
family. Incidentally, andesembiids and anisembiids
have no close relatives in the Old World; it is pos-
sible, however, that the rather enigmatic, Amazo-
nian genus Microembia Ross (Embiidae) might be
related.

It is likely that many additional species will be
found in the often-disconnected, difficult-to-reach
cloud forests and paramos of the Andes. Most of the
known species are placed in the highly diversified
new genus Andesembia. A strikingly-distinct spe-
cies is assigned to the monotypic new genus
Bryonembia. The two genera are distinguished, as
follows:

Andesembia: Adult males with left cercus two-
segmented. Anterior angles of clypeus not produced.
Size small, forewing length not exceeding 8 mm.
Known only from the Andean highlands of Ecuador
and Colombia.

Bryonembia: Adult males with segments of left
cercus fused. Anterior angles of clypeus conspicu-
ously lobed. Size large; forewing length exceeding
11 mm. Known only from a cloud forest locality in
central Peruvian Andes. Additional species should
occur in other Peruvian cloud forests.

Genus Andesembia Ross
new genus

Type species—Andesembia cuencae, new spe-
cies, by present designation.

Distribution.—South America: Highlands of
Ecuador and Colombia.

Diagnosis.—Males: Rather small; eyes usually
abruptly globose, often with pigmented facet-inter-
stices. Antennae with basal segment darker; flagel-
lar segments usually unicolorous to apex, but some
species have apical segments gradually pale; seg-
ments elongate, setae small. Mandibles usually very
large and highly irregular in form; however, in one
species (A. incompta) they are delicate, apically-ta-
pered, and minutely dentate. Submentum moderately
sclerotized; sides usually strongly divergent from
base to apex. Hind basitarsus very slender, excep-
tionally long, without a medial papilla; lateroventral
setae in basal half angled distad, the apicals pointed
ventrad. All wing veins behind RP represented only
by setae and well-defined pigment bands; MA un-
branched; cross-veins absent behind RP. Terminalia
with left hemitergite (10 L) small; its process
(10 LP) long, narrow, parallel-sided, usually curled
ventrad. Right hemitergite (10 R) without a promi-
nent process. Medial sclerite (MS) very large in all
species (except in 4. calinae) and fused to 10 R,
well separated by a membranous cleavage extended
to base of tenth tergite. Medial flap (MF) variable,
in some species almost obsolete. Epiproct sclerite
(EP) sclerotic, narrow, angled to the right beneath
MF and 10 R; its forward apex of two types: one
with a few large barbs which are angled to the right,
the other with a very dense patch of microspicules.
Ventral sclerite as described for the family, but with
right paraproct (RPPT) always a large, half-ring plate
pressed into the membranous, inner base of right
cercus. Left cercus always two-segmented; the basal
segment with a sclerotic outer-basal flange (which
may be a portion of a cercus-basipodite), inner side
of segment gradually expanded and increasingly
echinulate distad, but not forming an abrupt lobe.
Basal segment of right cercus weakly sclerotized;
inner-base deeply depressed by pressure of the
paraproct. Apical segments of both cerci normal,
equal.

Females.— Without noteworthy generic charac-
ters.

Biology.—Scattered, small colonies occur in
bark of tree stumps, as well as in draped moss and
lichen coverings on rock ledges in paramo zones and
damp cloud forests of higher altitudes of the Andes.
Usually colonies are very difficult to find because
of density of moss. In southern Colombia one spe-

ROSS:

cies, A. incompta, was found in deep moss growing
on tree fern trunks (Frontispiece), as well as under
bark flakes. The type species, A. cuencae, is com-
mon both in moss on large rocks and beneath bark
flakes of small trees.

The highest altitudinal record of any species of
the genus is 3700 meters, west of Cuenca in south-
ern Ecuador; the lowest is near Bafios, Ecuador, at
1700 meters. Cultures of all species are difficult to
maintain for they die within a few weeks after being
carried to lower, warmer altitudes. Collectors, there-
fore, are advised to gather study series of adults in
the field. Males in cultures usually matured during
February and March.

Component species.—Six new species were col-
lected from the Loja region of Ecuador in the south,
northward into mountains west of Cali, Colombia.
Because of the diversity of these species and the
often-disconnected geographic extent of suitable
habitats yet to be searched, the genus should com-
prise many additional species. Colonies are easily
overlooked in tangled, mossy habitats and invari-
ably consist of a single female and her brood. This
may explain why, in spite of concentrated search, I
have been unable, as yet, to find andesembiids in
cloud forest and paramo zones in Ecuador north of
Bafios. The habitats are suitable, species must be
present, but finding colonies is like “searching for a
needle in a haystack.”

Component species, all new, form three groups:
the Cuencae Group: two species from central Ecua-
dorian Andes, the Jncompta Group: one species from
extreme southern Colombia, and the Popayanae
Group: two species from western Colombia.

KEY TO SPECIES OF ANDESEMBIA
(Males)

1. Mandibles simple, similar to those of Oligotoma
and many other species of the order; molar por-
tion not strongly projected mesad. Cranium be-
neath antennal sockets not strongly or acutely
projected laterad. Southern Colombia.............

wiSesetuses ssvzeseissiencesseetacesieinvcaséossyseseshuct incompta

— Mandibles complex, inner faces deeply
emarginated; molar portion usually strongly
projected mesad. Cranium beneath antennal
sockets strongly projected laterad as acute or
blunt points. Ecuador and west-central Co-

EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 4 3

2. Epiproct (EP) with a few coarse, irregular barbs
at forward end which are angled meso-caudad.
ECUAG OLA rate terete tere ee 3

— EP with a conspicuous, dark, oval patch of
very dense microspicules on its forward end.
Golomb late iii acess cseeccvssttsspsssece cs tee ees 5

3. Outer basal angles of mandibles extensively and
very acutely projected laterad; dorso-basal sur-
face elevated as a narrow, broadly-angled carina.
Left tergal process (10 LP) straight.... banosae

— Outer basal angles of mandibles less projected,
apices blunt; dorso-basal surface without a
conspicuous carina. Left process (10 LP) curled
leftward and ventrad .............eceeeeeeeeeeeeee 4

4. 10 LP broadly expanded distad, only 2 barbs
on EP, caudal margin of LPPT narrower than
HP. Loja region, Ecuador... lojae

— 10 LP narrowly expanded distad, numerous
barbs on EP, caudal margin of LPPT as broad
AS Ee ee aeees ten reernees sees rere cuencae

5. Anterior angles of submentum narrowly pro-
jected forward. Popayan region, Colombia

sri Gesteae Mctee is cebeen oene eee aan eens popayanae
— Anterior angles of submentum broadly rounded.
Cali region, Colombia ......................... calinae

CUENCAE GROUP
Andesembia cuencae Ross
new species
(FIGURE 1)

Holotype.—Male, on slide, CAS. Data——Ecua-
dor: Lago Zhurucuchu, 3700 m elev. (est.), 11 km
W of Cuenca, Azuay, 16-II-1955 (E. S. Ross).

Description.—Appearance: Small (body length
7.5 mm); wings relatively large, extended well be-
yond apices of cerci; generally golden brown with
creamy white prothoracic membranes; cranium ma-
hogany brown; apical antennal segments only
slightly paler than others. Color details (in alcohol):
cranium dark mahogany brown except for paler ver-
tex pattern; eyes gray-black; antennal scape ma-
hogany brown, other segments, except three tan
apicals, medium brown (15 segments); preclypeal
membrane creamy white; mandibles dark amber with
piceous margins, other mouthparts medium brown
with creamy white membranes; submentum glossy,
piceous. Body sclerites and legs varied shades of
medium brown with creamy white membranes;

4 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 153

TERMINALIA
DORSAL

10 RP

TERMINALIA
VENTRAL

FIGURE |. Important characters of holotype of Andesembia cuencae Ross. Type locality: Ecuador, Lago Zhurucuchu,

Prov. Azuay (11 km W of Cuenca), est. 3700 m elev.

terminalia medium mahogany brown with piceous
margins, distal segments of cerci almost creamy
white in contrast to medium brown basal segments.
Dimensions (on slide): Body length 7.5 mm; fore-
wing length 5.5 mm, breadth 1.4 mm.

Important integumental characters—As figured.
Noteworthy is the shape of the mandibles without
acutely pointed outer angles, or a prominent dorso-
basal carina; submentum without forward projections
of the apical angles; cranial projections beneath an-
tennal sockets short, apically rounded; left tergal

process (10 LP) narrow, parallel-sided, not strongly
flared distad, curled ventrad; medial sclerite (MF)
broad, inner and caudal margins well defined; right
tergal process (10 RP) with a subapical membranous
“chevron” between medial flap (MF) and its outer
margin; inner side of basal segment of left cercus
(LC,) gradually expanded but not definitely lobed,
entire surface echinulate.

Paratypes.—A series of topotypic males depos-
ited in CAS, USNM, BMNH and NMQ.

ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 4 5

Allotype.—Female (in alcohol) with holotype
data and disposition.

Description.—Color details: Almost all sclerites
are dark chocolate brown; cranium, pronotum and
mesonotum darkest; first acrotergite pale yellow. All
membranes creamy white. Basal antennal segment
medium brown, all flagellar segments (18 in all) light
brown; apicals concolorous with basals. Cerci yel-
low tan with whitish membranes. Eighth abdominal
sternite evenly sclerotized except for two narrow,
pale excisions along caudal margin; color dark ma-
hogany brown. Ninth sternite also dark mahogany
brown; transversly narrow and broadly, shallowly
emarginated along anterior margin. Accessory gland
area membranous. Paraprocts light brown basally
blending to creamy white caudally. Body length: 8.5
mm.

Parallotypes —Series of topotypic females with
paratypic disposition.

Discussion.—The type cultures were first col-
lected by me (1955) and later by David Cavagnaro
(1964), in dense, low cloud forest near a small lake.
Colonies were common under bark flakes of trees
and stumps. More recently (1993) I found colonies
common beneath lichens and in moss tufts on large
volcanic boulders in open paramo of Parque
Nacional Cajas, 3550 m elev., about 20 km NW of
Cuenca. Because males matured during January,
February, June, and July, it is likely that maturity
isn’t seasonally limited.

David Cavagnaro also collected a culture of a
very closely-related new species, far to the south of
Cuenca in second-growth paramo scrub at 3000 m
elev. on the east slope of the misty pass between
Loja and Catamayo. Colonies were in crevices on
the bank of a deeply eroded Indian trail. Males
matured during July and August, 1964. The culture
did not long survive under laboratory conditions.
These specimens are treated, at least tentatively, as
a distinct species, described below.

Andesembia lojae Ross
new species
(FIGURE 2)
Holotype.—Male, on slide, CAS. Data—Ecua-
dor: Pass west of Loja, at 3000 m elev. on Loja-
Catamayo road (D. Q. Cavagnaro).

Description.—A ppearance, size and coloration

as described for A. cuencae. Integumental charac-
ters, as figured, showing minor distinctions, as fol-
lows: Projection beneath antennal socket narrowly
acute, slightly slanted caudad (blunt, projected
straight laterad in A. cuencae). Left process of tenth
tergite (10 LP) broadly flared (narrowly flared in A.
cuencae). Subapical surface of 10 RP solid (with a
chevron-shaped, membranous area in A. cuencae).
Epiproct sclerite (EP) with only two barbs (several
in A. cuencae). Caudal portion of LPPT narrower
than HP (broad in A. cuencae, as broad as HP).

Discussion.— I was tempted to regard the Loja
population as a subspecies of 4. cuencae; however,
with many intervening topographic barriers and dis-
tance, the two populations are significantly sepa-
rated. My attempt to collect specimens in interme-
diate localities failed—also failed in Podocarpus
National Park, a more southernly location. Although
difficult to find, colonies of Andesembia almost cer-
tainly occur in these regions.

Andesembia banosae Ross
new species
(FIGURE 3)

Holotype.—Male, on slide, CAS. Data——Ecua-
dor: 3 mi E of Bafios, 1700 m elev. Crevice in vol-
canic cliff, 11-II-55 (E. S. Ross).

Description.—Appearance: Small (body length
6.5 mm); wings rather small, short; body generally
dark brown, prothorax contrastingly pale. Color de-
tails (in alcohol): Cranium piceous dorsally and ven-
trally, lacking pattern. Basal antennal segment
concolorous with head, other segments medium
brown except the apical (15th), which is tan. Man-
dibles dark brown basally, blending distad to pale
amber, teeth reddish amber. Palpi medium brown,
submentum piceous. Prothorax straw yellow with
medium brown sclerites; forelegs dark brown ex-
cept for medium brown coxae and trochanters.
Pterothorax dark brown with slight bluish luster, legs
similar but somewhat paler. Abdomen reddish tan
with light brown sclerites; ninth and tenth somites
dark brown with paler cerci. Dimensions (on slide):
Body length 6.5 mm; forewing length 4.9 mm,
breadth 1.2 mm.

Important anatomical characters.—As figured.
Noteworthy are the extensively projected, acute,
outer angles of the mandibles, as well as the longi-
tudinal broadly angulate, dorsal carina of the

HEAD

OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 153

TERMINALIA
DORSAL

TERMINALIA
VENTRAL

FIGURE 2. Important characters of Andesembia lojae n. sp. Type locality: Ecuador, Loja-Catamayo road, crest of pass at

about 3000 m elev., Prov. Loja.

mandible’s basal third; the distally swollen anten-
nal scape; the straight, horizontal (not flared or
downwardly curved) left tergal process (10 LP); the
less coarsely-spiculated, forward end of the epiproct
(EP); the reduced left paraproct (LPPT); and the
short, globose basal segment (LC,) of the left cer-
cus which is rounded on its inner face and coarsely
echinulate on its entire length.

Paratypes —A small, topotypic series collected
by me (1955) and David Cavagnaro (1964), depos-
ited in CAS, USNM, and NMQ.

Discussion.—Colonies, consisting of one parent
female and brood, were rare and inconspicuous in
crevices in vertical, volcanic cliffs. They weren’t
found in silty trail banks, as was a small new spe-
cies of Oligembia, nor were they found in moss or
under bark flakes, as was the case with more abun-
dant 4. cuencae. David Cavagnaro’s small topotypic
culture produced five adult males during July, 1964,
but the culture soon died.

ROSS:

TERMINALIA
DORSAL

FIGURE 3 (A-C. Important characters of holotype of
Andesembia banosae Ross. Type locality: Ecuador, 3 mi
E of Bafios, Prov. Tungurahua, 1700 m elev.

EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 4 7

TERMINALIA
VENTRAL

INCOMPTA GROUP
Andesembia incompta Ross
new species
(FRONTISPIECE and FIGURE 4)

Holotype.—Male, on slide, CAS. Data——Colom-
bia: 25 mi W of Macoa, Prov. Putamayo, 2030 m
elev., 3-III-55, trunk of tree fern in paramo (E. S.
Ross).

Description.—A ppearance: Medium sized (body
length 8.0 mm); uniformly tan except for darker
brown head and prothorax; wings large, extending
beyond apices of cerci. Color details (in alcohol):
Cranium dark mahogany brown, pattern faint; eyes
gray-black; basal antennal segment medium brown,
flagellar segments tan, becoming pale to apex (18
segments); mandibles amber with piceous margins,
other mouthparts tan except for medium brown
submentum. Pterothorax and legs golden brown with
piceous margins and sutures; prothorax darker. Ab-
domen dark creamy white, becoming darker caudad;
terminalia concolorous with pterothorax, cerci tan.
Dimensions (on slide): Body length 8.0 mm; fore-
wing length 7.0 mm, breadth 2.0 mm.

Important integumental characters ——Most sig-
nificant is the narrow, elongate-oval cranium with
greatly reduced projections beneath the antennal
sockets; the small, simple mandibles with fine api-
cal dentation; the terminalia are basically similar to
those of species of the Cuencae Group, differing
primarily in reduced spiculation of the epiproct.

Paratypes.—Males from 2000-2500 meter zone
between Santiago and Macoa, Prov. Putamayo,

8 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 153

HEAD

TERMINALIA

TERMINALIA
DORSAL

VENTRAL

FIGURE 4. Important characters of holotype of Andesembia incompta Ross. Type locality: Colombia, 25 mi W of

Macoa, Prov. Putamayo, 2030 m elev.

southern Colombia. Deposited in CAS, USNM, and
ICNB.

Discussion.—It was almost disappointing to dis-
cover this species with its ordinary head and man-
dibles in a family otherwise characterized by unusual
head and mandible features. These simple charac-
ters alone are sufficient for immediate recognition
of the species.

The type series was collected in a few places
between Macoa and Santiago, Prov. Putamayo,
southern Colombia, between 2000 and 2500 m el-
evation. Habitats in the paramo zone include tree
fern epiphytes (Frontispiece) and mossy stumps in
cleared pasture.

ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 4 9

POPAYANAE GROUP
Andesembia popayanae Ross
new species
(FIGURE 5)

Holotype —Male, on slide, CAS. Data-—Colom-
bia: 11 mi N of Popayan, Prov. Cauca, 1830 m elev.,
5-III-55 (E. S. Ross).

Description.—A ppearance: Rather small (body
length 7.2 mm), wings large; medium brown
throughout with head piceous, antennal apices whit-
ish. Color details (in alcohol): Cranium piceous
throughout, without pattern, ventral surface paler
brown. Eyes subcutaneously gray-black. Basal an-
tennal segment concolorous with cranium, segment

HIND BASITARSUS

2 paler; segment 3 to apex blending from medium
brown to creamy white. Mandibles piceous basally,
blending to dark amber distad, dental margins red-
dish amber; palpi gray-tan, lacinia yellowish,
submentum piceous. Thorax and legs varied shades
of medium brown; prothorax darker; wings medium
brown. Abdomen paler than thorax; terminalia
scarcely darker, margins of sclerites and medial
sclerite (MS) piceous; basal segments of cerci light
brown, distals lighter. Dimensions (on slide): Body
length 7.2 mm; forewing length 6.0 mm, breadth 1.7
mm.

Important integumental characters —As figured;
noteworthy are irregularly indented, inner margins
of the mandibles; large, broad submentum with di-

10 RP

TERMINALIA
DORSAL

TERMINALIA
VENTRAL

FIGURE 5. Important characters of holotype of Andesembia popayanae Ross. Type locality: Colombia, 11 mi N of

Popayan, Prov. Cauca, 1830 m elev.

10 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 153

vergent sides terminating with pronounced projec-
tions of outer apical corners; left mandible with much
higher outer-basal ridge than on the right one; the
small, narrow, downwardly-twisted, left tergal pro-
cess (10 LP); the dark, densely-microspiculate, for-
ward patch of the epiproct; the strigose left paraproct
(LPPT); and the gradually distally-expanded, basal
segment of the left cercus which bears scattered
echinulations.

Discussion.—This species is readily recognized
by the very peculiar mandibles and submentum of
its males. Only the holotype and an accompanying
adult female were collected on a mossy tree base
projecting from aroad bank. The significance of the
species wasn’t realized until months later in my home
laboratory. Otherwise, the short road stop would
have been extended in the hope of obtaining a pro-
ductive culture.

The conspicuous dense patch of the epiproct’s
microspiculation relates A. popayanae to A. calinae,
but great differences in the mandibles, submentum
and other characters (as figured) readily distinguish
the two species.

Andesembia calinae Ross
new species
(FIGURE 6)

Holotype.—Male, on slide, CAS. Data——Colom-
bia: 13 mi W of Cali, Prov. Valle, 1820 m elev., 20-
III-55 (E. S. Ross).

Name basis.—Reference to type locality near
Cali.

Description —A ppearance: Medium sized (body
length 9.0 mm), wings large; medium brown
throughout, head piceous. Color details (in alcohol):
Cranium piceous without trace of pattern, ventral
surface paler brown. Basal segment of antenna dark
brown, second lighter, 3-7 smoke tan, 8—10 gray-
white, distal segments lost. Mandibles blending from
dark brown basally to amber subapically, dental
margins reddish amber; palpi medium brown;
submentum piceous, as dark, or darker, than cranium.
Entire thorax and legs varied shades of medium
brown, prothorax somewhat darker; mid- and hind
tarsi pale tan. Abdomen light brown with tan mem-
branes; terminalia, including basal cercus segments,
medium brown, apical segments paler. Dimensions
(on slide): Body length 9.0 mm; forewing length 6.6
mm, breadth 1.0 mm.

Important integumental characters.—As figured;
mandibles large, irregular in form, apices narrowly
tapered, inner bases strongly projected mesad;
submentum heavily sclerotized, but small, sides di-
vergent forward, apical corners rounded. Left tergal
process (10 LP) moderately large, parallel sided,
abruptly curved leftward, terminated as a sharp point;
right process (10 RP) very short. Epiproct (EP) with
a very dense, dark mass of microspiculations. Basal
segment of left cercus gradually swollen distad but
not forming a lobe, echinulations confined to distal
inner surface.

Discussion.—The type locality is on the road
between Cali and Buenaventura. A single colony was
found under bark of a damp, rotting stump in re-
cently cleared montane forest. The galleries deeply
penetrated the soft fibrous bark. Although numer-
ous nymphs were cultured, only the holotype and
nine females matured, all others died.

Genus Bryonembia Ross
new genus

Type species.—Bryonembia amplialata Ross,
new species, by present designation.

Name basis.—Greek bryon, moss, in reference
to the mossy habitat.

Distribution.—Peru: Cloud forest zone, Carpish
mountains (east of Huanuco).

Diagnosis.—Males: Very large, slender-bodied
with disproportionately-large wings but with abdo-
men and terminalia unusually small. Uniformly tan
with dark head, all appendages concolorous with
body. Cranium as in Andesembia, but with clypeus
medially depressed and with each anterior angle
greatly produced as a blunt lobe. Antennae with
flagellar segments very elongate. Mandibles broad-
based and angled outward, then abruptly angled
mesad at 90°; apical dentation normal. Submentum
narrower at base than at apex. Hind basitarsi as in
Andesembia, but with latero-ventral setae longer.
Wings also similar but proportionately larger.
Terminalia with basic structure of Andesembia, dif-
fering as follows: left tergal process (10 LP) shorter
and broader; right paraproct (RPPT) (or perhaps a
cercus-basipodite) smaller, fragmentary; left
paraproct (LPPT) without trace of apical
echinulations; distal segment of left cercus broadly
fused to basal segment, line of former articulation
marked by membrane.

ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 4

HEAD

10 LP
10 RP

TERMINALIA. “~~... f
VENTRAL nee

10L

TERMINALIA
DORSAL

FIGURE 6. Important characters of holotype of Andesembia calinae Ross. Type locality: Colombia, 13 mi W of Cali,

Prov. Valle, 1820 m elev.

Females.—Uniformly brown throughout, appar-
ently without generic level characters.

Biology.—Occurs in cloud forests which receive
very heavy rain or mist. The type species is most
readily collected on rocky road banks where the thin
silk galleries radiate from crevices. The species also
occurs in hanging moss on trees and vines, as well
as in trunks of tree ferns. The zone is about 3000 m
elevation. Adults of both sexes were collected in the
field during September and May and thus maturity
is not limited to one period each year.

Unfortunately, since my visit forty-eight years
ago, the habitat has probably been destroyed, or se-
riously damaged, by steep, hillside farming and tree-
cutting for fuel and charcoal production.

The silk galleries are very thin-walled and ex-
tend unbranched for considerable distances. Only
one species is known. The species does not survive
long in cultures and only late stage nymphs live long
enough to mature.

12 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 153

Bryonembia amplialata Ross Name basis.—Reference to exceptionally large
new species wings.
(FIGURE 7)

Description—As described in the generic de-
Holotype.—Male, on slide, CAS. Data.—Peru: _ scription, and illustrated. Dimensions (on slide):

2 mi (by road) E of crest of Carpish mountains, —_ Body length 12.5 mm; forewing length 11.5 mm,

Huanuco, about 3000 m elev., 17-IX-1954 (E.S. breadth 2.75 mm.

Ross).

TERMINALIA
DORSAL

10 RE

RPPT 19 pp.
TERMINALIA
VENTRAL

FiGuRE 7. Important characters of holotype of Bryonembia amplialata Ross. Type locality: Peru, 2 mi E of crest of
Carpish Mtns, Prov. Huanuco, about 3000 m elev.

ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 4 13

Paratypes.—Six topotypic males deposited in
CAS, USNM, and MUSM.

Allotype-—Female (in alcohol) with holotype
data and disposition.

Description.—Appearance: All sclerotized por-
tions of body and legs dark chestnut brown. Cra-
nium rather elongate, basically light chestnut brown
but clouded in a definite vertex pattern with darker
brown. Antennal scape pale amber, tinged with
brown; flagellar segments (17 in all) pale brown,
distal segment yellowish. Acrotergites concolorous
with nota. Eighth abdominal sternite with a narrow
dark brown crescent on each side, otherwise exten-
sively pale tan. Ninth sternite dark brown with a

membranous, obtuse emargination in anterior mar-
gin. Paraprocts tan. Basal segment of cercus medium
brown, distals pale tan. Body length 12.5 mm.

Parallotypes.—Four topotypic females in CAS.

Discussion.—It is tantalizing to consider the
occurrence of many other remarkable congeners in
scattered, often disconnected cloud forests of the
Peruvian and Bolivian Andes.

It should be noted that a very active, elongate
species of Onychophora occurs in the damp, mossy
roadbank habitat of Brvonembia and might be one
of its predators.

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