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MEMOIRS Read
OF THE
AMERICAN ENTOMOLOGICAL SOCIETY
0 sont NUMBER 14

CLASSIFICATION OF THE

BLATTARIA

AS INDICATED BY

THEIR WINGS
(ORTHOPTERA)
BY
JOHN W. H. REHN

PUBLISHED BY THE AMERICAN ENTOMOLOGICAL SOCIETY
AT THE ACADEMY OF NATURAL SCIENCES
PHILADELPHIA
1951 This publication is sent to
estabiisn date of issue.
Please siamp date of re_aipt

and piace in your Library.
FES 7 1951

MEMOIRS
OF THE

AMERICAN ENTOMOLOGICAL SOCIETY

* NUMBER 14

CLASSIFICATION OF THE

BLATTARIA

AS INDICATED BY

THEIR WINGS
(ORTHOPTERA)
BY

JOHN W. H. REHN

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PUBLISHED BY THE AMERICAN ENTOMOLOGICAL SOCIETY
AT THE ACADEMY OF NATURAL SCIENCES

PHILADELPHIA
IQS 1
? This publication is sent to
establisn date of issue.
Please stamp date of receipt
and place in your Library,

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(Issued January 31, 1951.)

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TABLE,OR CONTENTS

LiEFOGNICWIOM” “ese eehasice ciao SiG cua ont CIRCLE MON RR SCT TCR Son rac rence crea ica aac Pore 1
Gharacterizationvor theyBlattania, ia. tcc tcl wie eels oereie eo are sens menses 4
Glacsiicavonvonthemblattaniaes ss. occise o<-a ck Sotetcine ole erat cee eens 4
PIMC carrvndeAMPeA TT GM. VplIa O'S ster sever etisne yay st ancy Ghavajso.2' eve cevsiuisitanae a ats) aoodaiig tenet Slonaiet leileravexsyo/% 8
EXP eanlaltl OMMONMONETINS) «04ers SA cisieuldd Sus se i neicyece a ee nelae sleltie teres 10
Bes aba AMPs ats ise cs eis Gode, Sic een sl ehe glee Oiu-w aru laeleiejale nis aed etetaiers 12
Comparative Table of Terms Used in the Discussion of the Wings of the
IBIHIANFTGY 5.6:0.0-07020.0-0. EEL OATES Ore ORTIOR SCONCE IE CENTER er Beier INCRE crea 13
Comparative Table of Terms Used in the Discussion of the Venation of
lemme Ss evista tg climes eyarsy ates ey sw sh in :teses tclMea nye Sasa 'sapaiterin irene ay sunire eaogememeieeovaetave ie 14
WWHDES — 6.0.06 2G G0.0 51g Oo SIDSOre I et CNET Renn NE eet ORS Re ie Pei I 19
FOGG — o.0's eG cad. lao oid 6 ONO N RI IR SIC RCC IRA tien aes ae CCRC ae 21
NVC Bi OCS mmPM NRE I csee ie Sa ce} yin cetea eye ayes Sashes duane meee <asererobemser ayers simap ie oe 22
SV¥SCMATS AMTEMEGMEME ooocooansdeno cde cuooNaGucD oon oC DS DUO DDOddKOE 23
Tentative Key to the Families of the Blattaria ........................ 23
Dissran of Permily IRSkierewhy goseescdadeesseoaoceooc50 x 0cndoooudS 23
Palkypheaeidae sc'coooccc cons evde pono gconsedpandoyoDO DUD AgoOBBIOBOpacue 25
Tentative Key to the Subfamilies of the Polyphagidae .................. 29
POlyPMGiIA® scooocasouogncsc0090d0d 0000 D00ad DGRDaDNUeODUDO0UR GOD 30
Pkg ovodcdecmorsnneppodes mie ons Ess ope oosoCNocogem ads 31
Condit % obcopcudadueeon ec toleati nn acmenoo eo op.c.o bin.cd coc mare 33
aT isciiit Te MMR Re NTI. U4, che ity cag Subic She auawnrecnhian ese cepammieratn ale Went te hvala foals 35
Bey Esnid Clutch mune ea eee aes cost oven ces cue cutee a enue uaMerewe ie: aura. aveuspausiiel- aieteltevey serve 36
TL atiiaGhieyl So. rey See ee eee Mace Meare mcenha ate oo eo Pls FUaicea Taree Mole ee 38
Compmsoalinn. ~~ ondooqetmsSueeenno obo cmos nunc aucoKc cmp covlodaoe 39
IG CEOUDNTAC coconnbupe oes sodooc tod vcnaeHoUbUnoso Coane CAgOS 39
Ewilayrriaplimee® ccdccongaodcddeccunbo 00 gosoeunoonuaceucacccDd00 40
DiplOMiECES oscocccocadoosboepoooccoo Dodou CHa ONGeEDodad cre donaueE 42
PRmOSHMGEe occccosnoodagudeeudceebouocs Ondo DDD UO Noo Ud OaDUOMOGaOaD0 44
TRING 6.5 docoro.e:606's BO IG Oe DIL OD TIO Cais Gra oO Eo OES Gecocno come CmOr air 47

MEM. AMER. ENT. soc., 14.

i TABLE OF CONTENTS

Tentative Key to the Subfamilies of the Blattidae ...................... 48
Bilatitimae: soils Saat cain ade tese temboeeraneces o cigaiie lene) eerie en ciisi areae Ue eee 50
INy.ctiborinae: | 2 ceivatten.goers cca onus OCI SR Cee Oe era eter 54

Nyctiborini jee decnety oratusie crave eoiouno tus oi ave elm oe ICR ere 56
Megaloblattinin | saccades Socnnne ce bess mentation Giese oko ereeie tela neeaenne By/
Epilampromds Complegnees ace eo eee eee cee ee eee onccoc. 58
Epilampninae Bit cacetea eaaciae sere anya denise cate tease tor wees one eens 59
Galolampmrini “Sa dishe sande iin east ercecncnte ete cae Se ieee eee ee 62
Oniscosominil Gaon ect acco ek ee rare ae cree 64
Perisphaerimt saciei ccm ceenscansevc perverse nue are ee acca eae earn oie: wana een 66
DIBXxbKe{O}D) oD Cb lacrya winters Maly Meanciaccets itl sharin RcEe eu PCreET mare mreraaaen ceo Geaeeo 68
Witopeltimitys cic ates ots cleans oneney oepe ee eeN eR ea Le re cea Rar Na creme pec 70
Epilamprinit, 4h ames cata cinee ere aoe aetna aera rae eee eee 72
Nauphoetini pina Geese cua Ce ene eats aaa eaters 74
Panchloninits 0 setae tea Soe cee tear a caoe tee eee 75

SD cloreVet i MmnnAm Mm niconicttn sricicn tins os Oa Hea Siac Baon Goaudie orig 010 77
Paranauphoetiniy aiamssecipec a acme reer reer cls ra eae aa ee 79
Phoraspidinin sjcciotl ety eis chorea rcnen eitoe cra Rina ele erent 81
Paratro pita nig aaosanrete oct eaten tater eae ore a ee eet ICICI 82
Brachycolimae: 2. ccc.cc oat woven oceanic acs ee oer oe nee 83
Blaberinae lease eee eee PR rie i Bic eek yO RENTS Se ceEMPECAN ep tar tech 3c 86
Pseudomopoid™ Complexe aeaeeneee eae eee ere eee nar eae 88
Pseudomopinae® ox c.ct/ tied tints cee ahe ai eee ne ae 89
Parcoblattiit .ccciicsio scan ey eterna cineca eat eR RE ee eR 93
Eustégastini. 4 ccmececsencos Seo ere Ceo nee eee 95
Euphyllodromini@ 4s cose coe ane oer ee ane 96
Tschriopteniin «do siewdwe mteeceel cra ce a ee rt See oreo heen eee 98
Euandroblattiny: 225.252 oy Scns ee ener a nee ene 99
Nieoblattellini@ c.03/5ev5c aor ere oC Ok ae oe 101
Pseudomopini oc¢.aiie sn cece een ene a aa eee 102
BUSttelimt «6 sks, cede tocome nee ohne Eee ele aCe aCe eee 104
Supellini: ac nentacines beh See ee eae OE aE ee eee 105
Symplocimt 2.565 eer waciccrcle eee Pre ome tee Rta cess clara cee 107
Balltitr 9. d53 0 dengan ae oasis patra See Ochna: Cree rie roe 109
Betobiimae schon atte sic dt oS Jeers hace re ee ee oo 110
i of 0) ob hc) Manan RN MEANS Ceara a Ioictis a n:o Gin ono ae CeO Gaara CC OG hro0 Hols 4 bic 113
Chorisoneurini’ 4...) eee ea ee eee 115
Plectopterini.. i123 cd coche Me etie C Cen eee 117
Anaplectinaé | 6.4. sie cachet cee eS Cre ee Eee 119
Ceuthobiinade: 00h Lee eB Ae ae ray. eG cen 121
Qulopterygidde: <0: ti foe eee 123
RELETENCES:, 23. ice.sie srehe en aentines aioe a ae Tae aE eT oP 126

MEMOIRS
OF THE

AMERICAN ENTOMOLOGICAL SOCIETY
NUMBER 14

CLASSIFICATION OF THE BLATTARIA AS
INDICATED BY THEIR WINGS
(ORTHOPTERA )

By Joun W. H. REHN
Professor of Biology, Marlboro College

Research Associate, Department of Insects
Academy of Natural Sciences of Philadelphia

INTRODUCTION

The Blattaria have long been recognized as one of the major groups
of the Orthoptera. However, the classification of the major com-
ponents of the group has not progressed adequately with the descrip-
tion of newly discovered forms. In addition, the wings of this group
have never been surveyed to see if they would furnish a guide to the
relationships of the various components. It is for these reasons that
the wings of representative genera have been examined for the present
study. These organs have been found to present features that are use-
ful for the classification of this group.

An active association with the orthopterists at the Academy of
Natural Sciences of Philadelphia has allowed me to examine the un-
usually representative collections of Orthoptera at that institution.
Even more important, however, has been the opportunity to be present
at, and in many instances instigate, discussions on the classification of

FEB 7 1951

2 CLASSIFICATION OF THE BLATTARIA

the orthopteroids. At such times, has been made available the results
of years of intimate contact and research in this group. Both the
unwritten but mentally matured ideas as well as the impressions and
suppositions have been aired. During such sessions an interest in the
broad classification of the complex was first developed and later stimu-
lated. While obtaining entomological training at Cornell University
the desire for a revision of the classification of the orthopteroids was
further encouraged.

During the last decade attempts have been made by a number of
workers to clarify and revise the classification of various portions of
the order,’ particularly the Acridoidea (1 & 2) and Gryllacrididae (3).
As certain other groups, such as the Mantiaria (4), did not appear to
to be in such complete turmoil, the Blattaria were chosen for study.
An additional impetus to the study of this group was the remarkable
distinction between the classification of the recent and fossil forms.
Although the present work does not attempt to integrate these com-
ponents it is hoped that after obtaining sufficient knowledge of the
existing forms a correlation may be attempted.

It is with this unique preparation that the analysis of representative
material and a revision of the classification is presented. It is well
known that the Philadelphia group is interested in the broad classifica-
tion of the Orthoptera. With immense collections available and the
feeling that there is more than enough for all interested, this study has
been undertaken.

Grateful acknowledgment of the suggestions, criticisms and opinions
and for the basic ideas for this study are due to Professor W. T. M.
Forbes. It is as a result of his personal direction and the interest
started by his paper on the axillary venation of insects (5), that the
present study has been completed. Also to Dr. V. S. L. Pate for his
unlimited helpful understanding, interest and suggestions which have
materially aided in all stages of the present work are due my grateful
thanks. My sincere thanks go to the late Morgan Hebard and to
J. A. G. Rehn of the Academy of Natural Sciences of Philadelphia for
their aid, opinions and criticisms and for their large part in making the
present study a reality.

1 Subsequent numbers in parenthesis throughout the text refer to numbered
references to literature which appear at the end of the paper.

JOHN W. H. REHN 3

To the authorities of the Academy of Natural Sciences of Phila-
delphia for placing the material for the present work at my disposal are
due my sincere thanks.

For the present study sixty-eight genera of Blattaria were critically
examined and are illustrated, while numerous additional genera were
surveyed. In practically all cases, male and female of the same species
were available for comparison and study of sexual dimorphism. In
most cases the genotypic species was examined. This was done when-
ever possible in order that any changes found necessary would not
necessitate the revision of existing generic concepts. In those cases
where the genotype has not been examined, it is believed and hoped,
that the form selected does not differ significantly from the typical one.
In certain cases, small to fair sized series of the species, usually from
the same locality, were examined in order to study variation. This
latter has been particularly helpful in ascertaining what features may
be considered reliable. In all cases, where it is possible, the person
originally determining the individual figured has been mentioned.

The selection of genera for this study was carefully conceived.
Consultation with other orthopterists at the Academy of Natural
Sciences of Philadelphia and with Dr. W. T. M. Forbes, as well as
personal knowledge, was utilized. An attempt was made to pick repre-
sentative forms from each of the known distinctive, or supposed dis-
tinctive, groups of the Blattaria. It is, therefore, believed that at least
most of the major entities have been represented in this study.

Two supposed distinctive groups have not been examined. These
are the Nocticolinae and Archiblattinae. Individuals of neither of these
groups have been available for study. In working from the published
information concerning them, a tentative placement has been made.
It would seem that the Nocticolinae are extremely specialized Blattidae.
Their cavernicolous habits have probably led to the reduction of the
tegmina and wings. Asa result, few if any important venational fea-
tures are retained. The Archiblattinae would seem to be related to
the Blattinae. The degree of relationship cannot be ascertained from
existing illustrations or discussions. The genus Archiblatta would
seem to represent a very primitive, archaic type.

MEM. AMER. ENT. Soc., 14.

4 CLASSIFICATION OF THE BLATTARIA

It is well recognized that a study of a single set of organs or a single
system will not give one sufficient information for a complete and
detailed classification. It is believed, however, that this study of the
wings will at least clarify certain points and it is here used to establish
a modified classification.

CHARACTERIZATION OF THE BLATTARIA

The Blattaria as a group constitute one of the most easily recog-
nizable assemblages of the Orthoptera or for that matter of any of the
insects. It seems advisable to treat them as a series, or as a super-
family, in the suborder Dictyoptera. By so grouping them with the
Mantiaria and with an indefinite number of fossil forms in one suborder
their basic relationship is indicated. The following characterization
should serve to distinguish the Blattaria from the other Orthopterous
insects.

Usually broadly oval and somewhat flattened in form; head free, usually
inflexed, nearly or completely concealed by the shield-like pronotum; mouth
directed posteriorly ; maxillae and labium of an omnivorous or carnivorous type,
former with galea slender, latter with both the glossae and paraglossae equally
developed; maxillary palpi with 5 articles, labial palpi with 3 articles; ocelli, if
present, normally two; antennae long, filamentous, annulate and multi-articulate ;
pronotum large, normally transverse, laterally expanded; tegmina and wings
variable, fully developed to completely absent; legs strong, the three pairs essen-
tially similar; coxae large, approximate, and not separated by a quadrate sternal
plate; femora with apical spurs usually present; tibiae usually spinose; tarsi
5-jointed; abdomen usually with ten visible tergites; cerci prominent, jointed;
males with styles which are usually visible.

Eggs covered with some type of a capsula (odtheca), although some forms
retain this within the body and are ovoviviparous.

Alimentary canal long and sinuous, crop large, gizzard with powerful masti-
catory armature; eight tubular enteric caeca, malpighian tubules in six groups.

THE CLASSIFICATION OF THE BLATTARIA

Many diverse classifications of the Orthoptera have been evolved
since Linnean time. The relative rank of the major divisions and the
methods of this subdivision and arrangement have been numerous and
for the most part far from satisfactory. The Blattaria always have
been recognized as one of the major divisions of the order. As such
a division, they have been given a rank varying from a genus in

JOHN W. H. REHN 5

Linnaeus’ classification to an order by such workers as Handlirsch.
This latter worker recognized in addition the order Protoblattoidea for
a great mass of fossil forms. Moreover, he grouped all of these cock-
roaches with the mantids into the ‘‘Unterklasse; Blattaeformia.’”’ No
general agreement has been reached upon the relative rank of this
complex, and its subdivisions. At first, groups were based on rather
careful work, but many have grown by accretion until they are no
longer natural entities. A brief resumé of their classification follows.

In the tenth edition of Linnaeus’ (6) Systema Naturae (1758), the
genus Blatta was proposed and, with other genera now considered to
be Orthoptera, was placed in the Coleoptera. In his later work, Twelfth
edition (7, 1767), these same insects were placed in the Hemiptera.
In 1773, De Geer (8) recognized the artificiality of this arrangement
and proposed, for what is now generally called the Orthoptera, the order
Dermaptera. Although this name is the oldest for the entire group,
subsequent restriction has limited it to the forms commonly known as
Earwigs.

Olivier (9), in 1789, applied the name “Orthoptéres” to most of
the then recognized genera. This system was followed by Lamarck
(10) in 1801.

In a classification devised by Leach (11), in 1815, the blattids were
removed from the other Orthoptera and he gave to them the name
Dictuoptera. This has been changed by later workers to Dictyoptera.

Still another grouping was made by Latreille (12), in 1817. At
this time, he divided the order into two sections placing Blatta, Mantis,
Phasma and Forficula in the Cursoria and the remainder of the Orthop-
tera in the Saltatoria. This classification with minor additions and
changes was often used for a long period.

Burmeister (13), in his Handbuch (1838), arranged this group in
twenty genera and a number of subgenera. In all, over one hundred
and twenty species were recognized. He erected a number of the
genera which later workers have used as a basis for higher groups.
Certain characters of the wings were used for this classification, but
venation as such, was considered as incidental except in one case. Ser-
ville (14), in 1839, in the classic work on the Orthoptera, recognized
ten genera, one subgenus and eighty species of Blattaria. Throughout
the following decades a considerable number of workers described

MEM. AMER. ENT. soc., 14.

6 CLASSIFICATION OF THE BLATTARIA

numerous forms. The next major works were those of Henri de Saus-
sure (15, 1862, 1864) in which well over fifty additional forms were
described, and wing venation was used rather extensively.

The classic work on the Blattaria is that of K. Brunner von
Wattenwyl (16, 1865). In this he recognized eleven tribes, over fifty-
six genera, a number of subgenera and one hundred seventy-eight
species. In most cases, these tribes represent groups which are still
utilized. However, their limits often have been modified and their
relative rank changed. This publication erected what might be termed
the first modern classification. In addition, it was one of the early
works to use wing venation for diagnosis and characterization. Even
to this day, due to its comprehensiveness, it is one of the most helpful
pieces of literature concerning the group.

Walker’s catalogue (17, 1868) unfortunately appeared before the
author could see Brunner’s Nouveau Systeme. In it, six hundred
ninety-six species are recognized and these are placed in a great many
families. The same author (18), in 1869, issued a supplement which
for the most part includes reference to the species described by Brunner,
but also describes a few new forms. Neither of these advanced the
classification of the group, but more often added confusion instead of
clarification.

During the next decade, the numerous works of Saussure (19)
added materially to our knowledge of this group. At this time, both
Saussure and Brunner (20) were actively working and their various
publications supplemented and amplified their own and others’ findings.

Kirby (21), in his synonymic catalogue of the Orthoptera (1904)
divided the Blattaria into sixteen subfamilies. These were essentially
those erected by Brunner with the addition of the Chorisoneurinae,
Nocticolinae, Archiblattinae and Plectopterinae. In earlier work, his
Polyphaginae has been termed the Heterogamidae. Also the Chori-
soneurinae and Oxyhaloinae had been considered as a unit. In this
catalogue two hundred thirty-seven genera were recognized and almost
two thousand species (1997).

After the turn of the century several classifications of the orthop-
teroids were proposed. Verhoeff (22, 1903) claimed that the mantids
and blattids constitute a separate order and termed it the Oothecaria.
In 1905 Navas (23) called the same complex the Dictyoptera. The

JOHN W. H. REHN 7

following year Krauss (24, 1906) considered the blattids to consti-
tute a separate order.

Handlirsch (25), in his classic work of 1903, raised the orthop-
teroid complex to two subclasses. In one of these, the Blattaeformia,
he recognized six orders, one of these being the Blattoidea. In his
later work (26, 1906-1908) these subclasses were reduced in rank
to superorders, but the ordinal rank was retained for the various
components.

At about the time these changes in the basic classifications were
taking place Shelford and other workers were revising some sections
of the Blattaria. Shelford’s (1907, 1908, 1910) revisions (27) of
various subfamilies in fascicles of the Genera Insectorum helped clarify
the picture. Shortly before this J. A. G. Rehn (28, 1903, 1904)
started working on the complex. Later workers such as Chopard (29,
LeoZ7etes) silanitsea (30; 1915, etc); Hebard (31, 1916; ete)) and
others added materially to our knowledge.

Karny (32), in 1921, attempted a revision of the major components
of the orthopteroids and endeavored to correlate the recent and fossil
forms. This paper on the classification of the Orthopteroidea while
instructive and interesting has not been followed by many workers.
In the portion dealing with the blattids most of the conclusions do not
seem to have been based on the examination of sufficient material.
However, this attempt led Handlirsch to revise his previous classifica-
tion. Thus, the classifications appearing in such works as Schroeder’s
(33, 1921) and Kukenthal’s (34, 1929) handbooks are in actuality a
revised Handlirschian classification.

In numerous textbooks and some entomological treatises, a number
of workers have followed the tendency to split the group into a large
number of families. For example, Brues and Melander (35, 1932) in
their classification of insects recognized twenty-four families in the
Blattaria.

On the other hand, most of the working orthopterists have been
content to consider these entities as subfamilies or tribes. In recent
years, a few have recognized some of the entities as being distinct
families. However, very few have recognized all of these groups as of
family rank.

MEM. AMER. ENT. sSoc., 14.

8 CLASSIFICATION OF THE BLATTARIA

In 1928, Forbes (36), in his study of the axillary venation of
insects, pointed out some of the major differences in members of this
complex.

In recent years, in addition to workers mentioned earlier, Bei-
Bienko (37, 1941), Princis (38, 1946), Bruijning (39, 1948) and
others have added materially to our knowledge of the group. The
number of genera and species described by these workers is large.

Practically all workers have placed the new forms described within
the classic subfamilies. Chopard and Hebard have redefined some of
these components. An example of this is Chopard’s (40) recognition
of the Nocticolinae and Hebard’s (41) division of the Ectobiinae and
Chorisoneurinae in his treatment of the Australian forms of these
subfamilies.

No recent attempt has been made to consider the classification of
the Blattaria from the world-wide point of view. Numerous faunal
works and generic revisions have been published. Rarely generic
groups have been revised. With the accretion of knowledge the picture
has become confused. It is interesting to note that while Kirby’s cata-
logue is the latest for the group, since that time at least two hundred
seventy genera and over fifteen hundred species of Blattaria have been
described.

A conservative estimate of the size of the group at present would
place it at four hundred fifty recognizable genera and somewhat more
than thirty-five hundred species. It is well known that in various por-
tions of the tropics and subtropics, which appear to offer optimum con-
ditions, the fauna is as yet very incompletely known. No detailed, or
for that matter well-founded, estimates on the number of existing forms
can be made.

TEGMINA AND WINGS

The Blattaria while being basically winged insects, have achieved
through various specializations, practically all possible conditions and
combinations from fully alate to completely apterous (42). However,
no cases are known in which the tegmina are reduced without a corre-
lated reduction in the wing. In all but the groups of Blattaria with
appendiculate fields, apterous and subapterous forms are known. In
some of the complexes these conditions are widespread. In connection

JOHN W. H. REHN 9

with this, it is interesting to note that while uniform reduction in
tegmina and wings often occurs in both sexes there are numerous
instances in which such reduction occurs only in the females.

The system of names used in this study is based on the Comstock-
Needham system of wing venation as amplified by Forbes. In addition
certain terms used by various orthopterists have been incorporated.
As a result of the study, certain limitations and amplifications of these
terms were found necessary. In order to clarify this situation the
following definitions are given.

In the Blattaria the venation is more richly branched than in the
Comstock hypothetical plan. In all cases the major veins may be
homologized but in some cases this is not true with the major branches.
Usually where the homology is not evident these branches have been
termed rami or sectors.

In some cases subcosta divides apically so that the branches are
apparently comparable to Sc: and Scz but in forms where the branches
are numerous no homology can be determined.

The division of radius into Ri and a radial sector is usually evident,
but the numerous subsidiary branches do not appear to have a homo-
logue in the Comstock-Needham system. The apical posterior branch
of the radial sector may actually represent the R1+5 of their system,
but sufficient evidence is not available to draw definite conclusions.

In media it seems probable that the major branches when present
are homologous with those of the hypothetical plan.

The cubitus with its great number of branches in most forms again
differs from the hypothetical type. However, in those groups where
there is a primary subdivision into stems or major branches, these may
well be homologous to the Cu, and Cu, of the Comstock-Needham
system.

The first plical vein, as pointed out by Forbes, is equivalent to the
vein Cu, of some Neuropterists and to the Ist A of others. It probably
represents the CuP of other workers and is the vena dividens of orthop-
terists. It is not included in the Comstock hypothetical plan.

The second plical vein is called 1st A in the hypothetical form.

The third plical vein, which is present only in the wing, is not noted
in the hypothetical wing plan.

MEM. AMER. ENT. Soc., 14.

10 CLASSIFICATION OF THE BLATTARIA

Posterior to the convex fold, are the anal veins. At least the first
of these is branched, and has been termed the branched axillary. The
anal area is much more richly veined than in most other insects and
the possible homologies with the anal veins of the hypothetical type
have not been determined.

Explanation of Terms
(Figures 1, 2, 3, 4, 5)
Alar Units

Tegmen. In the usual orthopterist style, used throughout for the mesotho-
racic wing.
Wing. Restricted to the metathoracic wing.

Margins

Anterior margin. Used for the costal margin of the tegmen to the point
where it rounds or passes into the apical margin. In the wing it is used for the
same relative portion or until it is intercepted by the fold which delimits the
appendiculate field.

Apical margin. Used in the tegmen for the margin of the apex from the
end of the anterior margin to the point where it passes into the posterior margin.
In the wing it is used for that portion of the margin from the end of the an-
terior margin to the plical notch; or in those forms having an appendiculate
field for the margin of this field. When a well-developed appendiculate field is
present, it and its margin are usually divided into an anterior and posterior
portion.

Posterior margin. Used in the tegmen for the portion of the margin from
the apical margin to the plical notch. This is, in part, the sutural margin of
some authors. This term is not used in the wings except for a portion of the
margin of the appendiculate field.

Margin of anal field. In the tegmen this is the margin of the tegmen basad
of the plical notch. In the wing, this is the margin of the wing posterior to the
plical area or in those forms with an appendiculate field the portion posterior
to its base.

Areas or Fields

Humeral area. Used in the tegmen for that portion of the tegmen anterior
to the subcosta. It has not been found necessary to consider it as a separate
entity in the wing.

Plical field. The area bounded anteriorly by the first plical and posteriorly
by the third plical vein.

Intercalated triangle. The folded portion of the plical field which in repose
is usually triangular in shape. In the Pseudomopinae it is normally bounded
anteriorly by the first plical and posteriorly by the third plical vein.

JOHN W. H. REHN 11

Appendiculate field. That portion of the wing which is set off by a trans-
verse fold; found in the Diplopteridae, Ectobiinae, Anaplectinae and Ouloptery-
gidae.

Veins

Costa. The vein forming the anterior margin of the tegmen and wing,
usually extending for a varying distance along the apical margin.

Subcosta. In the tegmen and wing this is the first vein behind the anterior
or costal margin. It may be simple, forked apically, with anterior rami, or with
posterior branches.

Anterior rami of subcosta. These are the branches going forward toward
the costal margin before the tip of the vein. They may be regular or not, simple
or branched; rarely clubbed in the wing.

Posterior branch of subcosta. Used for a condition noted in the tegmen.
Apparently the basal anterior rami of the radius have become secondarily
stalked on the subcosta and resemble a posterior branch.

Radius. The second major vein behind the anterior margin. It usually
divides into a distinct R, and a radial sector.

R,. This vein usually becomes free from the remainder of radius near the
base. Anterior rami may be present or not. When some of these rami have
become secondarily stalked on the subcosta they are termed the posterior branch
of subcosta. At times a pseudostigma is formed in the wing along the anterior
rami.

Anterior rami of R,. These are similar to the anterior rami of the subcosta.

Radial sector. Used for the remainder of radius after the branching into R,.
It may have anterior and apical sectors as well as the apical posterior branches.

Anterior rami of radius, or anterior rami of radial sector. These are the
branches that go toward the anterior margin, comparable to rami of the sub-
costa and R,. They may be simple, irregular, secondarily branched, or fused
apically.

Apical ram of radius, or apical rami of radial sector. These are the anterior
branches of the radial system that curve forward and toward the apical margin.
They may be simple, or secondarily divided. Terminal twigging may be present.

Apical posterior branches of radius, or apical posterior branch of radial
sector. Used for the most posterior main branch of the radial system possibly
equalling R45. It usually terminates at the apical margin, but may have
secondary divisions which in turn may be branched. Usually, it is on a line
with the main stem of radius, but may be posterior to this.

Media. This is the third main vein behind the anterior margin. It may be
simple or have a number of branches, which may further subdivide. In general,
a tendency towards the simplification of this vein is found throughout the group.
Its origin is intimately connected, as a rule, with the arculus, usually rising from
the top or middle of this vein. Rarely, it may be stalked basally on the radius.

Cubitus. This is the fourth vein stem behind the anterior margin. In the
tegmen, it divides into a number of branches (sectors). At times, there are

MEM. AMER. ENT. soc., 14.

12 CLASSIFICATION OF THE BLATTARIA

two major divisions each with further subdivisions. These primary subdivisions
are termed stems or major branches. In the wing, the branches are normally
termed sectors and they may go to the apical margin or posteriorly to the plical
furrow. All branches and sectors may show secondary division.

First plical vein. This is a vein immediately behind the cubitus and before
the plical fold. It is usually a simple vein extending to the margin. In the
tegmen, it apparently lies at the base of the plical furrow. In the wing, it is
before the plical fold or furrow and rarely has terminal branches. This vein
is apparently equivalent to the CuP of many authors.

Second plical vein. This vein is not evident in the tegmen except as a
trachea. In the wing, it is immediately behind the first plical vein and béfore
the plical fold. Primitively it is complete, passing to the wing margin, how-
ever, it is often reduced. It may be free apically or its apex may curve forward
and join the first plical vein. At times, its basal portion is completely lost.
This vein is apparently the second A of most of the holometabolous insects and
the first A of the neuropteroids.

Third plical vein. This vein is not recognizable in the tegmen. In the
wing, it is usually prominent, lying behind the second plical and when free,
rising from the same sclerite, and before the branched axillary. It usually
passes to the wing margin behind the plical notch when that is present. In
forms with an appendiculate field supported by venation, this vein forms the
support. Basally, it may be free or it may be stalked on the branched axillary.

Both the first and third plical vein may show some apical branching or
twigging. This is particularly true in the Polyphagidae.

Branched axillary (Branched anal). In the wing this vein rises from the
basal bar and is immediately behind the third plical vein. Typically, it gives
rise to a variable number of branches, which are usually simple and further
apart apically, but rarely there may be secondary branches. At times its more
‘posterior branches cannot be differentiated from the remaining veins in the anal
field.

Anal veins. In the tegmen all the veins posterior to the plical furrow are
indicated by this term. In the wing these are the simple veins posterior to the
branched axillary.

Arculus. In part an oblique or vertical crossvein and a portion of media,
evident only in the wing. It is a strong, simple vein connecting the bases of
media and cubitus. When media apparently rises from its middle, it may appar-
ently connect radius with cubitus.

Tegmina
(Figure 1A and B)
The tegmina (mesothoracic wings) vary considerably in texture,

form, venation and apparently in function. Primitively their function
seems to be not only an integral part of the flight mechanism, but also

JOHN WwW.

H, REHN

13

COMPARATIVE TABLE OF TERMS USED IN DISCUSSION OF THE
WINGS OF THE BLATTARIA

Margins &
Areas

Anterior mar-
gin (tegmen)

Anterior mar-
gin (wing)

Apical margin
(tegmen)

Apical margin
(wing)

Posterior mar-
gin (tegmen)

Margin of
anal field
(tegmen)

Margin of
anal field
(wing)

Humeral area

Plical field or
intercalated
triangle

Appendiculate
field

Hebard (45)

Margin of
marginal &
scapular field
(pt.)

Margin of an-
terior field
(pt.)

Margin of
scapular (pt.)
& of discoidal
(pt.) fields

Margin of an-
terior field
(pt.)

Margin of dis-
coidal field
(pt.)

Margin of
anal field

Margin of pos-
terior field

Area bounded
by marginal
field

Intercalated

triangle

Appendiculate
field

Shelford (46)

Margin of
mediastinal
and marginal
(pt.) areas

Margin of
marginal (pt.)
& discoidal
(pt.) areas

Margin of dis-
coidal area
(pt.)

Margin of
anal field

Mediastinal
area

Triangular
apical area

Saussure &
Zehntner (47)

Margin of
basal area
and of mar-
ginal field

Margin of an-
terior field
(pt.)

Margin of
marginal field
(pt.) and dis-
coidal area
(pt.)

Margin of an-
terior field
(pt.)

Margin of dis-
coidal field
(pt.)

Margin of
anal field

Margin of
posterior, in-
termediate or
axillary and
radiate fields

Basal area

Saussure (48)

Margin of
mediastine &
costal (pt.)
areas and of
marginal field

Margin of
marginal field
(pt.)

Margin of
costal area
(pt. ?), mar-
ginal (pt.)
and discoidal
(pt.) fields

Margin of
marginal (pt.)
and discoidal
(pt.) fields

Margin of dis-
coidal field
(pt.)

Margin of
anal field

Margin of
anal (pt.) and-
posterior
fields

Mediastinal
area

Intercalated
or reflexed
field

Brunner (49)

Margin of
marginal or
mediastine &
scapular field
(pt. ?)

Margin of
scapular field
(pt.)

Margin of
anal field

Margin of
anal field
(pt).

Marginal or
mediastinal
field

Triangular
apical field

In this study, fields,
supporting the area, i.e.

except as noted above, are named for the vein
radial field, median field and cubital field.

MEM. AMER. ENT. soc., 14.

OF THE BLATTARIA

CLASSIFICATION

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MEM. AMER. ENT. Soc., 14.

16 CLASSIFICATION OF THE BLATTARIA

a covering for the metathoracic wings when these are in repose, and a
protection for the abdomen. In the more advanced forms their use in
flight is apparently more and more restricted while the protective value
is amplified. In groups such as those with an appendiculate field they
appear to serve mainly as a protection for the wing and abdomen. In
females of some of the polyphagoid genera, which have relatively soft
bodies and reduced wings, the tegmina apparently protect the dorsum
of the abdomen.

In some of the groups of the Blattaria, the tegminal texture is
essentially similar to that of the wings, but in most they are at least
somewhat thickened, leathery or coriaceous. From this a further ad-
vance in thickness is noted particularly in the Corydiini, Perisphaerini
and Oulopterygidae. In these the venation while still discernible is not
prominent. An extreme condition is to be found in Diploptera in
which the venation is subobsolete, the vein disposition being indicated
by a few tracheae and the clear areas between punctures. The portion
of the one tegmen that is covered when in repose is usually more mem-
branous, or at least less coriaceous, than the tegmina as a whole. In
all material studied, this has been the right tegmen. Rarely, as in
Holocompsa, the tegmina are thickened and leathery basally and mem-
branous apically.

In shape the tegmina, while basically elongate ovoid, vary from
practically subtrigonal on the one hand to subrectangulate or subquad-
rate on the other. In most instances these latter conditions are corre-
lated with varying degrees of wing reduction. The subtrigonal shape
and its modifications occur when a distinctive form is necessary to
cover the folded hind wing, and where the tegmina act only as a pro-
tective cover for the abdomen.

The tegmina are normally richly veined and in general have the
following characteristics. The costa is strong and bounds the anterior
margin. Possibly in some forms there is basally a trace of a precostal
area. The subcosta is practically always strong, either simple, or with
a varying number of anterior rami; at times it appears that at least one
of the radial rami become stalked upon this vein. A ventral develop-
ment of subcosta into a flange-like flap is noticed in some groups and
it would seem to act as a tegminal catch or lock. The radius is also
a strong vein and usually occupies much of the anterior portion of the

JOHN W. H. REHN 17

ANAL —
PIOSTEM CON
1A eS as

POST.

MAR. ANAL | POST. MAR.

Text-figure 1.—A and B, diagram of tegmen.

tegmina. The apical posterior sector, while prominent in many of the
more primitive forms, is progressively shortened and apparently lost in
some of the more highly developed groups. The development of a
number of anterior rami of the radius would appear to be one of the
distinctive features of the Blattaria tegmina. Media, in the more
primitive groups, is a strong, moderately branched vein but becomes
reduced to a single stem. It may become either stalked basally on
radius or cubitus. The marked reduction of this vein with the corre-
sponding expansion of either radius or cubitus or both is again charac-

MEM. AMER. ENT. sSoc., 14.

18 CLASSIFICATION OF THE BLATTARIA

Text-figures 2 to 5.—Fig. 2, diagram of wing. Fig. 3A, diagram of wing
folding in Blattidae. Fig. 3B, cross section of 3A. Fig. 4A and B, diagram of
wing folding of form with an appendiculate field. Fig. 4C, cross section of 4B.
Fig. 5, diagram of wing folding in Polyphagidae.

Explanation of Abbreviations

ANT. MAR., anterior margin; AP. MAR., apical margin; MAR. ANAL,
margin of anal field; POST. MAR., posterior margin; C, costa; CU, cubitus;
H, humeral area; M, media; R1, radius 1; SC, subcosta; ANT. R. RAMI, an-
terior radial rami; ANT. RAMI SC, anterior rami of subcosta; AP. RAMI,
apical radial rami; AP. POST. BR., apical posterior branch of radius; BR. AX.,
branched axillary; PL. FR., plical furrow; STEM CU, stem of cubitus; 1 PL.,
first plical vein; 2 PL., second plical vein; 3 PL., third plical vein.

teristic. Cubitus often divides into two main stems. The more pos-
terior of these stems usually fills, with the exception of the well-marked
anal area, the posterior half of the tegmina. The anterior stem forks

JOHN W. H. REHN 19

and may further subdivide. In the lower groups these divisions usually
terminate at the posterior margin, but in the higher forms a reduction
in the number of branches is noticed as is a change in their direction.
The paralleling of the posterior margin by these branches and their
ending at or near the apex is characteristic. The posterior branch of
cubitus (CuP) lies in the distinct plical furrow and appears to be
homologous with the first plical vein of the wing.

The plical furrow marks off the anal area and reaches the posterior
margin at varying points in different groups. Whether the second
and third plical veins are present in this fold has not been determined.
A trachea representing the second plical is shown in Comstock’s Wings
of Insects (50) behind the furrow.

The area behind the anal furrow constitutes what has been retained
of the anal fan. While the course of several veins may be traced, their
actual homologies are not known, but tracheation indicates that both
an anterior and posterior fan are present.

Wings
(Figures 2-5)

The wings (metathoracie wings) while less variable in function
and texture are considerably more diverse in venational types and
form. When they are developed flight would appear to be their only
function and when they are reduced so that this is not possible they
appear to be functionless. They are usually membranous, rarely
somewhat thickened and subcoriaceous. Exceptionally pseudostigmal
areas are developed.

In form, wings that are definitely capable of supporting flight may
be divided into two main types, first the polyphagoid with its large
preaxillary and small anal areas, and second, that found in other exist-
ing types with a relatively large preaxillary area and with a greatly
enlarged anal area. In the former, the length of the axillary furrow
is always less than one-half the total wing length, usually about one-
third, while in the latter this furrow is nearly equal in length to the
total wing length except in those forms with an appendiculate field.
In the groups with an appendiculate field this field is developed from
the interplical area except in Diploptera and Oulopteryx, and it may
be equal to twice the length of the radius. In Diploptera all major

MEM. AMER. ENT. soc., 14.

20 CLASSIFICATION OF THE BLATTARIA

venation crosses the transverse fold, thus representing still another
type of development. In Oulopteryx an appendiculate field without
venation is present. How this has arisen and its homologies are
not known.

The venation of the wings is extremely diverse. Most of the
major groups of the Blattaria have their own peculiar type of speciali-
zation. The costa forms the anterior margin of the wing and extends
for a variable distance along the apical margin. In general, the sub-
costa is relatively prominent and may be either simple, forked or
have rami, often with terminal twigging. The anterior rami are at
times clubbed. At times capture of the anterior rami of radius is
noted. When this happens they have been termed the posterior
branches of the subcosta.

The radius is always well detec and normally has a distinct
Ri, which may be simple or have anterior rami as in the subcosta.
In some cases, it is not distinguishable from the anterior rami or
the remainder of the radius. Normally the remainder of the radius
has a number of anterior and apical rami, these latter often secondarily
dividing. Usually an apical posterior branch of radius is present.
This may be either simple or show secondary division. At times
it has become indistinguishable or lost.

The media is always retained but varies from a primitive type with
several distinct branches, to a simple, single vein. This vein may
be taken as a guide to the degree of specialization of the group under
consideration. The media usually apparently rises from the arculus,
either from its top or from the middle; however, it may be stalked
on the radius. The arculus is present in all but some of the Panes-
thiidae and is normally strong, either vertical or oblique.

The cubitus shows many types of specialization. Primitively it is
a major vein with numerous diagonal, oblique sectors. One line of
specialization is that having a division into at least two main stems
each showing further subdivision. In some groups these sectors tend
to pass to the plical furrow instead of to the apical margin. This
tendency is carried to its extreme in certain of the Epilamprinae. On
the other hand, a tendency towards the simplification and reduction
in the number of its branches is evident in the Pseudomopinae. In
this group the number of branches tends to be reduced and all attain

JOHN W. H. REHN ail

the apical margin. In forms such as those with an intercalated field
it may be reduced to a simple, single stem.

The first plical vein is normally well developed and complete. In
many of the Polyphagidae it has some terminal branches or twigging.
However, in the Anaplectinae only the apical portion of this vein is
retained the base being lost. This is the only group in which this
condition has been noted. The second plical vein shows considerable
variation. In the more primitive types it is often complete, extending
to the apical margin. In different groups it is reduced to varying
degrees. In these assemblages all conditions from free and complete
to very short and ending in the first plical vein can be found. The
third plical vein is normally complete. It is primitively free from the
branched axillary in such groups as the Blattinae, Polyphagidae and
others. In this latter group it often has apical branching. In more
specialized groups it may be stalked on the branched axillary to vary-
ing degrees. This condition is noted particularly in the Epilamprinae
and is carried to an extreme in the Thoracini. Another type of
development is found in the Ectobiinae and Anaplectinae; in these
forms the third plical has been elongated so that it is the only support
for the appendiculate field. In the Diplopteridae the third plical
supports the posterior half of the appendiculate field and has many
irregular branches. The anterior portion of the field in this genus
is supported by the major venation which crosses the transverse fold.

The branched axillary vein normally has two or more branches
and may have as many as nine or ten. These are usually simple
branches, but rarely some secondary branching may be noted. The
area supported by this vein may be rather limited or may be equal
to one-half of the anal fan. In the Polyphagidae there may be three
branched axillaries. Behind this are a number of simple anal veins.

Crossveins are normally present throughout most of the wings,
and it is only in certain of the more specialized forms such as the
Ectobiinae, Anaplectinae and Oulopterygidae that they are reduced to
a number that can be used in classification.

Folding.—The tegmina are not capable of being folded.

The wings have a number of methods of folding, but may be
considered to consist of three basic types. First, the polyphagid type
with its definite anterior and posterior fan in the anal area. The

MEM. AMER. ENT. Soc., 14.

22. CLASSIFICATION OF THE BLATTARIA

wing folds along the plical area separating the preaxillary from the
anal area. In this type the anal area is not folded fanwise. In all
of the other groups the wing folds along this same area, but the anal
area is folded fanwise, with alternating convex and concave folds.

Those forms with an appendiculate field, and the Diplopteridae and
Oulopterygidae all have the fold along the plical area and a fan-like
folding of the anal area, but, in addition, have the apical portion
of the wing reflexed. This fold may only be slightly indicated with
the apical area crumpled as in Ectobius or may be distinctly developed.
The fold may be chevron-shaped or transverse. In these forms this
apical area is normally reflexed on the dorsal surface of the wing.
However, in the Oulopterygidae this area is coiled into a tight spiral.
Thus not only by venation but also by the method of folding are the
major groups of the complex more or less indicated.

Methods

In this study it was found necessary, in practically all cases, to
remove the wings from the specimens examined. The large anal fan
and the basal portion of the plical area can only rarely be examined
by any other means. Usually both tegmina and wings were floated
onto glass in either water or weak alcohol. By adjustment and
combination of transmitted and reflected light details could be deter-
mined. In a few cases balsam mounts were prepared but were found
less satisfactory than the wet preparations. The long established
practice of orthopterists of mounting wings on cards, or linen ledger
is definitely unsatisfactory as it is only through the use of transmitted
light that some features can be ascertained.

For the drawing, the wings were often projected so that major
features and proportions could be obtained. After this the sketch was
checked, against the projected wing and the opposite wing of the same
individual, and in most cases against additional material. No attempt
was made to keep the drawings to any definite scale as the extreme
variation in size would have resulted in drawings too small to be useful
on the one hand and so large as to be cumbersome on the other.

After finishing with the studying of the wings, they may be fastened
in their original position on the specimen, kept in fluid, made into
permanent slides or into a permanent mount that can be placed be-
neath the specimen (51).

JOHN W. H. REHN 23

SYSTEMATIC ARRANGEMENT

The existing Blattaria are divisible on venational features into
two large complexes, possible superfamilies. The first includes only
the Polyphagidae while the other includes the remainder of the Blattaria
centering upon the family Blattidae, but including the Panesthiidae,
Diplopteridae and Oulopterygidae. The interrelationship and phylo-
genetic placement of some of these entities is not clearly evident. The
Nocticolinae and Archiblattinae have not been included, as material of
these two groups was not available for study.

Tentative Key to the Families of the Blattaria (Alate Forms Only)

1. Wings folding only along the plical fold .................. POLYPHAGIDAE
Wings with anal area plaited in repose as well as folded along plical fold .. 2
2. Wings with apical portion marked off by a transverse fold
Wings without a distinct apical portion marked off by a transverse fold .. 5
3. Apical portion of wings without venation, coiled in a spiral in repose.

OULOPTERYGIDAE

Apical portion of wings with some venation, either folded or crumpled in

LE POSE MEM GVietwC Oil Cie stere eee yaseataredene ersraesanece ia) ecacduei euskal eomcampeleuatevWravevensees Sen 4

4. All major elements of venation crossing transverse fold of wings; tegmina
elytriform, their venation subobsolete ................. DIPLOPTERIDAE

Only a single plical vein in apical portion of wings; tegmina normal and
WIN WSRAWIOMN GWGSME ooconcccvcg0g000000bN0uDNCeE BLATTIDAE (part)

5. Wings with five or fewer simple branches in radial system, no anterior or
apicaliramieyanculus oftentabsent ann.) 4+ scmeee ee dae PANESTHIIDAE

Wings with more than five branches in radial system, anterior or apical rami

or both present; arculus always present ............ BLATTIDAE (part)
OULOPTERYGIDAE

BLATT IDAE
DIPLOPTERIDAE

PANESTHIDAE
POLYPHAGIDAE

ANCESTRAL STOCK

Diagram 1.—Probable relationship of the families of the Blattaria.

MEM. AMER. ENT. Soc., 14.

24 CLASSIFICATION OF THE BLATTARIA

The following arrangement appears to represent the most logical
linear sequence of the groups recognized in this study. However,
like all linear arrangements it does not present a true picture of the
relationship of the complexes.

The generic names used for the formation of group names are not
always the oldest used for the group involved. However, they have
been chosen because of their lack of ambiguity and in the belief that
they are representative of the assemblages involved.

Family Subfamily Tribe

Polyphagidae Polyphaginae Polyphagini
Corydini
Tiviinae Tiviini
Holocompsinae Holocompsini
Euthyrrhaphinae Euthyrrhaphini
Latindiinae Latindiini
Compsodini
Panesthiidae Panesthiinae Panesthiini
Diplopteridae Diplopterinae Diplopterini
Blattidae Blattinae Blattini
Nyctiborinae Nyctiborini
Megaloblattini
Epilamprinae Calolamprini
Oniscosomin1
Perisphaerini
Leucophaeini
Litopeltiini
Epilamprini
Nauphoetini
Panchlorini
Thoracini
Paranauphoetini
Phoraspidini
Paratropini
Brachycolinae Brachycolini
Blaberinae Blaberini
Pseudomopinae Parcoblattini
Eustegastini
Euphyllodromiini
Ischnopterini
Euandroblattini
Neoblattellini
Pseudomopini
Blattellini
Supellini
Symplocini
Baltini
Ectobiinae Ectobiini
Chorisoneurini
Plectopterini
Anaplectinae Anaplectini
Ceuthobiinae Ceuthobiini
Oulopterygidae Oulopteryginae Oulopterygini

bo
oun

JOHN W. HB. REHN

POLY PHAGIDAE

The Polyphagidae have the tegmina less distinctive than the wings,
but they possess a number of features of considerable diagnostic value.
The presence of numerous subcostal and anterior radial rami is a
condition parallel to that found in the primitive members of the
Blattidae. The cubitus can be considered as a rather primitive type
when compared with conditions found in other groups. <A_ well-
branched media, more freely branching than in other forms, can be
considered both distinctive and as basic to the type from which the
other Blattaria were developed. The finding of the first plical in the
base of the plical furrow, and in at least some instances with its basal
connection to cubitus still intact, shows the relationship of these veins.
This condition is more primitive than that in the more evolved blattids.
The occasional posterior development of the radius denotes a type of
specialization not occurring elsewhere. In many of the members of the
family the tegmina are membranous and very flexible. However, in
the specialized forms, they have become densely coriaceous, more so
than in practically any other groups except the Diplopteridae.

The wings are so distinctive in shape that they are immediately
separable from all the other members of the Blattaria. The large pre-
axillary area and the small unfolded lobate anal fan are unique. This
area may have some wrinkles or incipient folds in the jugal area. The
presence of a series of branched axillary veins, as well as the three
complete plical veins, some of which show terminal twigging, separate
the members of this family from all the others. The retention of a
branched media is in common with other primitive forms of the
Blattaria. The oblique cubital sectors are reminiscent of conditions
found in the Blattidae, particularly in the Blattinae, and these con-
ditions might be taken as basic to those found in such groups as the
Epilamprinae. The presence of a well-developed radial system is also
suggestive of that found in the primitive Blattidae.

In general the tegmina are elongate oval, either transparent and
membranous or densely coriaceous. The anterior margin is weakly
to strongly arcuate, with its greatest anterior expansion being approxi-
mately where the subcosta reaches this margin. The apical margin
is strongly arcuate to subobtuse angulate, roundly passing into the
posterior margin which is regularly arcuate to the plical furrow. This

MEM. AMER. ENT. soc., 14.

26 CLASSIFICATION OF THE BLATTARIA

latter is marked, as a rule, by a slight indentation or notch. The
margin of the anal field is rounded to the tegminal base. The humeral
area, particularly in the primitive forms, may be expanded, as in some
Epilamprinae, but in the more specialized subfamilies this area shows
reduction.

The subcosta is primitively heavy basally, straight to arcuate and
with numerous anterior rami; rarely in certain advanced forms with
posterior branches (Dyscologamia). This vein becomes simpler with
advancement and in the most highly evolved forms is simple and almost
straight. In this, it parallels the condition found in the Blattidae,
while in the Panesthiidae, Diplopteridae and Oulopterygidae this vein
is also heavy and simple, sinuate or arcuate. This vein normally
reaches the anterior margin in the basal one-third to one-half in those
forms with membranous tegmina. In those with coriaceous tegmina,
which are normally somewhat reduced, it reaches this margin at or
beyond the middle. The radius is usually well developed, but does
not have a distinct Ri. Primitively, it has a considerable number of
crowded anterior rami and a distinct separation between these and
the apical rami. In the advanced types there is less crowding and
better differentiation, resembling the condition found in the Blattidae.
To some extent the primitive condition is reminiscent of that found
in the Panesthiidae. The radial sector primitively has a number of
apical rami and posteriorly directed branches. These latter shift
forward in higher forms and pass to the apex, comparable to the con-
dition found in the basic Blattinae and Blaberinae. The freely branch-
ing media primitively has three or more posteriorly directed branches,
but soon these pass to the apical margin as in the other Blattaria. In
the higher Polyphagidae it is represented by a single branched vein,
always distinct from the cubitus as in some of the epilamproids.
Cubitus may show a division into two stems as in many of the Blattidae
or may be reduced. This vein shows various developments in some of
the subfamilies. Its sectors are at least arcuate and in general go
to the posterio-apical angle as in the Blattinae, but do not tend to
parallel the posterior margin as in many of the higher Blattaria. The
plical furrow is well indicated, definitely angulate in the primitive
forms, resembling the condition found in the Perisphaerini. The more

JOHN W. H. REHN Dd

advanced types have it evenly arcuate as in most of the Blattidae ex-
cept the pseudomopoids. The strength of the arcuation is in part
characteristic of some of the subdivisions. In at least some cases, the
first plical (CuP) is evident in the furrow and its basal connection
with cubitus can be ascertained. However, this is not true of the more
specialized forms. The anal area approximates in length the subcosta
as in the Blattidae, not longer as in the Panesthiidae. The anal veins
show more evidence of branching in the anterior portion of this field
than in most forms. In general, the veins curve toward the apical
corner as in the Nyctiborinae and Blaberinae.

The wings have a distinctive shape, the large preaxillary area and
small unfolded anal lobe which will separate them from all other Blat-
taria. Also, the short plical fold, usually one-half or less of the wing
length, as opposed to being almost equal to the wing length in all
the other forms. The anterior margin varies from straight through
weakly arcuate to weakly biconvex. The apical margin is either
broadly or narrowly rounded and passes into the arcuate posterior
margin. <A definite and decided notch is always noticeable at the
margin of the plical fold. The margin of the anal field is usually
regularly and evenly arcuate.

The subcosta which is relatively long but not distinctive may have
anterior rami, twigging or not. This vein reaches the anterior margin
beyond the basal third, often near the middle. Ri is longer and as a
rule more nearly simple than in most of the Blattidae, much as in the
Panesthiidae. In the primitive forms some indefinite, irregular an-
terior branches are present, while in some of the more specialized
forms this vein is simple. In certain of the subsidiary groups rather
definite and regular branching is found. The radial sector has both
anterior and apical rami in the primitive genera and these are more
distinct than in other Blattaria. Some advanced types approach con-
ditions found in Blattidae. Specialization is carried on by (1) a re-
duction or loss in the number of anterior rami so that only the apical
branches and branches of the radial sector are retained; (2) a reduc-
tion in the branches of the sector and a regular distribution of anterior
rami; (3) the loss of the apical branches. Rarely some of the branches
of the radial sector extend arcuato-sinuately to the wing apex. How-
ever, this seems to be merely a generic characteristic. The arculus is

MEM. AMER. ENT. soc., 14.

28 CLASSIFICATION OF THE BLATTARIA

strong, prominent and transverse. Media may apparently rise either
from the arculus or be stalked on the radial system. In the former
case, its origin may be at the top or in the middle of the arculus. The
mode of origin of this vein is helpful in defining the subsidiary groups
within this family. The media is strong, primitively with three or
more definite and some indefinite basal accessory branches. In special-
ized forms, the accessory branches are lost and the number of major
branches may be reduced to two; or in the extreme case of Holocompsa
the media is an incomplete stub. The cubitus has a large and variable
number of diagonal sectors all going toward, and most attaining, the
apical margin, a condition not found elsewhere in the Blattaria. It
occupies a larger area than any other system. An archedictyon is
present at the base of cubitus as in the Brachycolinae and certain other
primitive genera. The first plical vein primitively has terminal
branches, but is simple in higher forms, in the latter being comparable
to the condition found in other Blattaria. The second plical vein is
normally simple and strong as in certain Nyctiborinae and Blaberinae.
Rarely, it is sinuate and with adventitious branches. The third plical
vein is well removed from the branched axillary as in the Blattinae,
Nyctiborinae and pseudomopoids. At times, intercalated veins are
found in the plical area. The first branched axillary has more sub-
divisions than in other Blattaria. Often in the primitive forms a
second and third branched axillary are present, a condition not found
in other Blattaria. Behind this, there are usually four or more simple
anal veins, rarely branched, which join the basal bar, but are distinct
from the attachment of the branched axillaries as in the Blattinae,
Nyctiborinae and in certain other forms.

Although there is extreme sexual dimorphism in many of the mem-
bers of this family, the females in those genera which possess wings
show essentially the same characters as do the males except that in
certain cases the tegmina have become thickened and leathery. In
some genera such as Corydia this condition has been attained by both
sexes. However, in a number of genera the females are completely
apterous.

Diagram 2.—Probable relationship of the components of the Polyphagidae.?
From checking a series of other than wing characters during this survey, the
same conclusions can be attained. Therefore, it is believed that this grouping
is warranted and logical.

2In the charts attempting to show relationship, a single line ( ) is used
between subfamilies and a double line ( ) between tribes and their subfamily.

~

ie)

JOHN W. H. REHN 29

The family may be considered to have the following components :
Polyphaginae, with the tribes Polyphagini and Corydiini.
Tiviinae, with only the nominate group.

Latindiinae, with the tribes Latindiini and Compsodini.
Holocompsinae, with only the typical group.
Euthyrrhaphinae, with only the distinctive genus Euthyrrhapha.

Tentative Key to the Subfamulies of the Polyphagidae
(Alate Forms Only)

Practically no venation visible in tegmina; wings with a pseudostigma

EUTHYRRHAPHINAE

Venation evident in at least basal portion of tegmina; wings without a
PSCUGOSE OITA tect enser rai tele caren oe Mie w sustenijsusvel thaieantocnale ers Gieee s 2

No venation in apical portion of tegmina; wings with R, and media clubbed.
HOLOCOMPSINAE

Venation throughout tegmina; wings without any clubbed veins ......... 3
Subcosta of tegmina with anterior rami; wings with more than one branched
Axa Lip Vary Pattee iG UArctic ewahiss HS oops ests @ outtess Coamsye aia a RAs PoLYPHAGINAE

Subcosta of tegmina without rami; wings with only one branched axillary .. 4
Radius of tegmina with regular apical branches; R, of wings with regular

APICAMDLANCMeSmemt tre cuatro oes ysikte cis oe ome niaere moeie oe TIVIINAE

Radius of tegmina with at most a short simple apical posterior branch; R, of

WAS Sine See Oe yA ccacbaccauccosvcccocducs LATINDIINAE
LATINDIINI COMPSODINI

Ros

HOLOCOMPSINAE LATINDIINAE

gee
ay CORYDIINI

2. ae

ANCESTRAL STOCK

Explanation of diagram on opposite page.

MEM. AMER. ENT. soc., 14.

30 CLASSIFICATION OF THE BLATTARIA

POLYPHAGINAE

The species of the typical subfamily, Polyphaginae, are all relatively
large forms and may be distinguished from members of the other
subfamilies by a combination of features. The tegmina have a
moderate to ample humeral area resembling the Tiviinae, but not the
other subfamilies, and the subcosta has at least some anterior rami
while the related groups lack these. The anterior rami of Ri are not
regularly spaced as in the other subfamilies and show some secondary
branching; the remainder of the radial system is full and has numer-
ous branches. It does not show any definite signs of reduction as in
all others in the family. The media is strong, well developed, with
three or more branches. Cubitus is rather full, much as in the
Tivinae, and the first plical vein (CuP) either shows a definite basal
connection with the remainder of cubitus or approximates it. The
plical furrow is relatively strong and distinctly arcuate.

The wings are relatively large, membranous and with the typical
form for the family. They do not show any signs of specialization by
reduction or simplification as do the wings of the other subfamilies,
but retain an abundant, richly branched venation. Those genera
showing reduction in wing size have retained the fundamental features
of venation characteristic of this group. The anterior wing margin,
except in those forms showing reduction, is relatively straight as in
most members of the family. The subcosta may show some signs of
twigging, but is usually simple. The first branch of radius may be
simple, as in the Compsodini, or show a number of anterior rami that
are generally irregular and incomplete. However, in the genus Hetero-
gamia they are more regular, but never evenly spaced as in the
Tiviinae. The radial sector is variable either with some anterior irreg-
ular rami and apical rami or with apical and posterior branches;
always well developed but not appearing crowded toward anterior
margin as in other subfamilies. Media is distinctive with three or
more usually rather irregular branches except in those forms having
reduced wings. The cubitus has a larger number of branches than
in related forms, which vary from regular to rather irregular; some
twigging usually is present in the basal area. The first plical vein
normally has at least two apical branches and at times it 1s even
further subdivided. This vein is simple in related forms. The third

JOHN W. H. REHN 31

plical vein is usually complete, but occasionally is rather indefinite
and irregular. Some intercalated veins are often present in this area.
The first branched axillary vein normally has a considerable number
of branches which may be regular, although more often they are rather
irregular or indefinite. In addition, there is at least one other branched
vein in the anal fan, often two or more. In other subfamilies only one
branched axillary is present, but in the Tiviinae terminal twigging
is present in what would appear to be one of the other axillaries. In
at least some cases, all the veins in the anal fan show definite branching.

This subfamily may be divided into two readily recognizable tribes:
Polyphagini and Corydiini.

Polyphagini

In the Polyphagini the females are apterous, while in the Corydiini
they are alate.

The males have a combination of features that substantiate the
differences found in the females. The tegmina have the humeral area
less developed and the rami of the subcosta are usually more irregular
and more crowded than in the Corydiini. At least the apical posterior
branches of radius, and usually other sectors as well, are directed
apico-posteriorly instead of passing directly to the apical margin or
curving anteriorly as in the Corydiini. Media often has its sectors
sinuately, instead of arcuately, passing towards the apico-posterior
angle; its base is often connected by branches anastomosing with
cubitus instead of being free. The cubitus shows at least a slight indi-
cation of an arcuation or angulate curve away from the plical furrow,
instead of more or less paralleling it as in the Corydiini. The join-
ing of the first plical vein (CuP) with cubitus is evident in some
members of the complex and has not been noted elsewhere.

In the wing, Ri usually shows some anterior rami. The radial
sector may have posteriorly directed sectors or more numerous apical
branches than the Corydiini. Media is variable from relatively simple
as in the Corydiini, to multibranched, but rises from the top of the
arculus. The cubitus is inclined to show more basal twigging than
in the other tribe. The anal area usually has branched veins posterior
to the branched axillaries, a condition not found in other groups
except the Corydiini.

MEM. AMER. ENT. Soc., 14.

Be CLASSIFICATION OF THE BLATTARIA

The tegmina of the male are always fully developed and mem-
branous, at times thickened. The subcosta is sometimes thickened
and usually has a number of irregular anterior rami. The radius
normally has a number of irregular anterior rami, and a number of
posterio-apically directed rami from the radial sector; some of these
latter having secondary branches. The media is full and has four
or more sectors which pass arcuately and usually sinuately toward the
posterio-apical margin. The cubitus is well developed and usually
has some incomplete anterior basal branches, often curving distinctly
away from the plical furrow, so that an isolated sub-crescentic area
is left without major venation. Usually a considerable number of
more or less irregular branches are present, while in some cases the
cubitus divides into two main stems each having secondary divisions.
In at least some instances CuP is directly connected with cubitus,
while in others the two are more or less approximate. The plical
furrow is generally arcuate. An indefinite and irregular number of
anal veins are present, some of these, particularly those in the anterior
portion, show branching.

Wings of normal shape, and fully veined. The subcosta is rela-
tively strong and either simple or with some anterior rami. Ri has
anterior rami, either regular or irregular. The remainder of the
radius has a number of apical sectors, these either irregular and with
secondary branching or regular; rarely somewhat sinuate. Media
rising from the top of the arculus, often with basal anterior adventi-
tious branches some of which join the radial system. Media with at
least three and usually more well-defined branches. Cubitus well
developed, with a number of diagonal sectors, some showing second-
ary branching, in all cases with considerable twigging in the basal
region. First plical vein strong and with some terminal branching.
Second plical vein either strong and regular or with adventitious
branches. Third plical vein usually strong and regular, but occasion-
ally irregular. First branched axillary with a number of major
branches, usually these showing some subdivision. Usually the
second axillary shows some branching and the third axillary always
shows distinct branching. The remainder of the anal veins may be
either simple or branched.

JOHN W. H. REHN 335)

Representatives figured: Arenivaga bolliana (Saussure) figs. 6,
76. ([Det. Hebard, 1937.] This species is the genotype. Male.
Brownsville, Texas.

Eupolyphaga sinensis (Walker), figs. 13, 77. [Det. Hebard.]
This species is the genotype. Male. Ching-yang-fu, Kansu.

Heterogamodes roseni (Brancsik) figs. 7, 82. [Det. Chopard. ]
The genotype is Heterogamia ursina Burmeister, [== Heterogamodes
ursina (Burmeister) ]. Male. Buiram-Ali, Transcaspia. U.S.S.R.

Polyphaga aegyptiaca (Linnaeus) figs. 14, 81. [Det. Hebard,
1932.] This species is the genotype. Male. Zakaki, Cyprus.

Corydiini

In this tribe which can only be compared with the Polyphagini,
both sexes have at least the tegmina preserved; usually the wings
are present, but they may be considerably reduced.

The tegmina vary from the normal for the group to somewhat
reduced, obovate and densely coriaceous. This latter condition is
first noticed in females, but is apparent in both sexes of certain genera.
The humeral area is more developed than in the Polyphagini, and in
those forms with coriaceous tegmina may be broadly expanded. The
subcostal rami are usually more regular than in the other tribe and are
never as crowded. The radius does not have posteriorly directed
branches, most of them going to the anterior margin and some to the
apical one, instead of curving posteriorly. Media has its base free
and its branching is regular and direct as opposed to the conditions
found in the Polyphagini. The cubitus does not curve as distinctly
away from the plical furrow and CuP is never found joined to cubitus.

In the wing both subcosta and Ri are usually simple instead of
with anterior rami as in the Polyphagini. In the radius the sectors
go to the apex, instead of posteriorly, and usually have but little
secondary branching. Media is either stalked on the radius or rises
from the middle of the arculus as opposed to the condition found in
the other tribe. Cubitus usually has less basal twigging than in the
other tribe. There are always some simple anal veins.

In those forms with fully developed tegmina, the humeral area is
moderately developed and the subcosta is usually strong and possesses
a number of anterior rami. In forms showing reduction the humeral

MEM. AMER. ENT. soc., 14.

34 CLASSIFICATION OF THE BLATTARIA

area is broader and anterior rami, while present, are less strongly
developed. A number of anterior rami are present in the radial
system, these being longer and usually more regular than in the other
tribe. Some of them show secondary branching. Apical rami are
present, but no posterior branches are developed in this tribe. The
media is well developed having at least four major branches, these
often further subdividing. Cubitus is more or less regularly rounded
along the plical furrow, not showing the sub-crescentic venation-free
area. It possesses a number of branches, but does not show a basal
division into two main stems. The plical furrow is regular and
more or less evenly arcuate, not approximating the base of cubitus.
In the more primitive forms, a number of anal veins are present;
however, in the more specialized forms most of these are lost and
only traces of two or three of these veins may be found.

The wings may be either reduced or not; those reduced in size
have a corresponding reduction in venation. The subcosta is long,
simple and straight, usually not reaching the costal margin. Ri is
usually prominent, simple and not attaining the costal margin. The
remainder of radius has some anterior and apical rami that are more
or less irregular. No posterior rami are present. Media may be
stalked basally with radius or rise from the middle of the arculus.
It does not show any basal adventitious branches and has at most
three relatively short branches with terminal twigging. In forms with
reduced wings, it shows simplification. The cubitus has a number
of sectors, usually more regular than in the other tribe. In the more
primitive forms some basal twigging is noticed, but in the more ad-
vanced genera this tendency is lost. The first plical vein shows
terminal twigging in the fully alate genera, but is simple in those
showing reduction; the second and third plical veins are simple, strong
and direct. In the fully alate forms, there are three branched axillary
veins, while those showing reduction have at least one branched axillary,
but the remainder may be reduced. There are always two or more
simple anal veins.

Representatives figured: Corydia petiveriana (Linnaeus) fig. 8.
[Det. Hebard.] This is the genotype. Male. Trichinopoly, Presi-
dency Madras, India.

~

JOHN W. H. REHN 35

Dyscologamia pilosa (Walker) figs. 15, 16, 83, 87. [Det. Hebard,
1928.] The genotype is Dyscologamia cesticulata Saussure. Male.
Grot by Baso, Sumatra. Female. Fort de Kock, Sumatra.

Homoeoganua mexicana (Burmeister) figs. 9,78. [Det. Hebard.]
This is the genotype. Male. Jalapa, V. C. Mexico.

TIVIINAE

This subfamily occupies an intermediate position between the
generalized Polyphaginae and some of the more specialized members
of the family. It might be characterized as showing a simplification
by reduction of the basic type of venation, but not showing any other
definite specialization.

The tegmina have a simple subcosta as in the advanced sub-
families, but the remainder of the tegmina are essentially like those
of the Polyphaginae. The anterior radial rami are numerous as in
the latter, but show the regular spacing of the Latindiinae. The
radius has apical branches as in the Polyphaginae, but they are more
regular than in that group. The media is sinuate as in the Poly-
phaginae, but has fewer branches, only two, and has not become as
simplified as in the Latindiinae. Cubitus has a number of oblique
branches, this again showing simplification from the primitive type,
but not as advanced as the most specialized groups.

The wing shape is like that of the Polyphaginae. The subcosta
is simple as in the Corydiini, but also is fused with the base of radius
as in the Euthyrrhaphinae. Ri has regular anterior rami not as in
the other tribes and the remainder of the radial system has simple
apical branches, an intermediate condition. Media is two branched
as in the Compsodini. The cubitus has rather regular oblique sectors,
an intermediate condition for the family, and lacks the basal twigging
found in most of the primitive forms. A\ll the plical veins are simple
as in the advanced subfamilies, and only one branched axillary is
present.

The tegmina have a simple, but heavy subcosta. The radius has
a number of regular anterior and apical rami and the apical posterior
radial branch is present. Media, which at first closely approximates
radius becomes free and has two complete simple branches. The
cubitus has a number of diagonal regular sectors. The plical furrow
is evenly arcuate. No veins can be found in the anal area.

MEM. AMER. ENT. soc., 14.

36 CLASSIFICATION OF THE BLATTARIA

The wing has a simple, direct subcosta and a four branched Rx,
with its branches very regular and direct. The radial sector has a
number of spaced, regular branches. Media rises from the middle,
or a little above the middle, of the arculus and has two simple, direct
branches. The cubitus has a number of regular sectors. All of the
plical veins are present, complete and relatively strong, the first and
second being somewhat sinuate. Between the third plical and the
branched axillary is a series of intercalated veins. The branched
axillary is moderately developed and has four main branches. The
remainder of the anal veins are simple except for a slight tendency
to terminal twigging.

From this complex apparently has been derived the three special-
ized subfamilies, Latindiinae, Holocompsinae and Euthyrrhaphinae.

Only the genus Tivia is placed in this subfamily.

Representative figured: Tivia termes (Karny) figs. 10, 85. [Det.
J. A. G. Rehn, 1933.] The genotype is Tivia simulatrix Walker.
Male. Vryburg, British Bechuanaland, South Africa.

LATINDIINAE

This subfamily consists of a specialized assemblage that has
apparently developed from the Tiviinae. The extreme specialization
of the tegmina reminds one of the condition found elsewhere only in
Ceuthobia, but this resemblance is purely superficial. The wing is
definitely polyphagoid.

The tegmina may be compared with the Tiviinae from which they
differ by having radius with anterior rami and at most a short simple
apical posterior branch instead of having regular apical branches and
a long apical posterior branch. Media more nearly parallels the
radius, than in the other group and the cubitus has fewer and more
regularly spaced branches which go to the posterior margin instead
of to the apical margin and apico-posterior angle. The plical furrow
is less arcuate than in the Tiviinae. These forms have the subcosta
and radius basad much as in the Holocompsinae, but the lack of a
complete vein system separates this latter group.

The wings are best compared with those of the Tiviinae, but
show some relationship to the Euthyrrhaphinae. The subcosta 1s
simple as in these groups, but differs by not being stalked on the
radius. RR: is either simple or bifurcate, thus differing from the con-

JOHN W. H. REHN 37

dition found in either of these groups, and the remainder of radius
has only anterior rami, another distinctive feature. The media is
either two branched as in the Tiviinae or simple. The cubitus has
fewer, at most five, regularly spaced, arcuate branches; none are
secondarily branched as in the other groups. The plical veins are
simple as in all the higher Polyphagidae. The axillary is either simple
or with one branch, a condition not found elsewhere in the family
except in the highly modified Holocompsinae.

The tegmina, while of normal shape and membranous texture,
do not resemble those of any of the other subfamilies. The subcosta is
simple, straight and unbranched. The radius is but moderately well
developed having a number of more or less regular, simple anterior
rami. The radial sector is either greatly reduced and not branched,
or not differentiated. The media is relatively strong and has two or
three simple branches, these ending at the tegminal apex. The cubi-
tus is somewhat reduced, the branches are moderately regular and vary
from three to ten; secondary branching is found. The plical furrow
is indicated, but is less strongly arcuate than in most members of the
family. The anal field either lacks evident venation or has only three
or four unbranched veins. An irregular network of large cells is
made by the crossveins giving the tegmina an entirely different appear-
ance from that of the other members of the family, and most other
Blattaria.

The wings have the characteristic shape for the family and show
specialization by a reduction in the venation, however, this is not as
extreme as in the Holocompsinae. The subcosta is simple, free and
direct. Rs: is either simple and direct or with an apical branch. The
radial sector has a fairly large number of anterior rami, rarely some
of these showing banching; no true apical or apical posterior sectors
are present. The media rises at the top of the arculus and is simple,
showing at most a division into two main branches. The cubitus
is prominent, but has only four or five simple, spaced, arcuate sectors.
The first plical vein is free, complete and direct. The second plical
vein is complete and from weakly to strongly sigmoid. The third
plical vein is free and complete. The first axillary vein may have
one major branch or may be simple. When it has any branches,
secondary twigging may also be present.

MEM. AMER. ENT. soc.. 14.

38 CLASSIFICATION OF THE BLATTARIA

This subfamily may be divided into the two tribes, Latindiini and
Compsodini. These two specialized groups although evidently rising
from a common type are readily separated. In the Latindiini, the
tegmina have the subcosta free from the radial system, media is stalked
basally on cubitus and shows but two branches, the cubital fan is rather
well developed, and no veins are evident in anal fan. On the other
hand, the Compsodini have the radius stalked on subcosta, media,
which has three branches, free basally from cubitus, the cubital
fan is somewhat reduced, and three anal veins are evident. The wings
of the Latindiini have a simple media,a more fully developed cubitus, a
less strongly sigmoid second plical, and they retain a branched axillary.
The wings of the Compsodini retain a branched media, the cubital fan
is more reduced; and the second plical vein is strongly sinuate. No
branching is evident in the axillary area.

Latindiini

The tegmina of this group have the normal shape for the family,
but have highly specialized venation. The subcosta is simple, strong,
and free. The radius has a number of more or less regularly spaced
anterior rami, and the radial sector is not evident. Basally, media
is stalked on cubitus and after becoming free passes to beyond the
apical fifth before dividing into two simple branches. The cubitus
after becoming free from media shows seven or eight more or less
regular sectors. The plical furrow is evident, lowly arcuate. No
veins are found in the anal field.

The wings show considerable reduction. The subcosta is simple
and somewhat curved. The radius has a rather well developed Ri
with one branch and the remainder of radius has a few weak anterior
rami; no apical posterior sector is evident. Media rises from the
top of the arculus and is a simple vein extending practically to the
wing apex. Cubitus occupies most of the pre-axillary area and has
five spaced, simple, arcuate sectors. The first plical vein is free and
slightly curved; the second plicai vein is complete and more strongly
sigmoid; the third plical vein is simple and direct. The first axillary
shows some branching; posterior to this there is only one simple
anal vein.

In this tribe is placed only the typical genus.

JOHN W. H. REHN 39

Representative figured: Latindia dohrniana Saussure and Zehntner
figs. 17, 80. [Det. Hebard, 1919.] Latindia maurella Stal is the
genotype. Female. Trinidad River, Panama.

Compsodini

The tegmina have the usual form for the family, but show a high
degree of specialization. Subcosta is heavy and is stalked basally
with the radius; after becoming free, the latter, has a few anterior
rami, and there is a short, simple, apical posterior sector. Media is
relatively strong and in the apical half divides into three branches that
go to the apex. Cubitus is but moderately developed and shows
three oblique sectors. The plical furrow is evident and arcuate. The
anal field has four simple, irregular veins.

The wing has a simple, direct subcosta. R: is well developed and
simple, and the remainder of radius has some sinuate anterior rami,
apically one of these sub-dividing. No apical posterior sector is
retained. Media rises at the top of the arculus and somewhat beyond
the middle of the wing divides into two simple branches. The cubitus
is moderately well developed with four simple, spaced, arcuate sectors.
The first plical vein is straight, direct and complete; the second plical
vein is simple and direct. The third plical vein is simple and direct.
All of the veins posterior to this are simple and incomplete basally.
No branched axillary is present.

Only the genus Compsodes is placed in this tribe.

Representative figured: Compsodes delicatulus (Saussure and
Zehntner) figs. 18, 84. [Det. Hebard, 1916.] This species is the
genotype. Male. Cacao, Trece Aguas, Guatemala.

HoOLOCOMPSINAE

This subfamily represents an extreme development of the Poly-
phagidae and does not appear to be closely related to any other known
group.

The lack of any venation in most of the apical portion of the teg-
mina is a unique condition. The basal portion with its simple subcosta
and radius with rather regular anterior branches resembles the con-
dition found in the Tiviinae and Latindiinae. Media and cubitus are
apparently fused basally and are incomplete, again a distinctive feature.
The plical furrow is definitely angulate as in some Polyphagini, while

MEM. AMER. ENT. soc., 14.

40 CLASSIFICATION OF THE BLATTARIA

the lack of venation in the anal field is a condition found in some other
members of the family.

The wings are unique with their large apical expansion of Ri and
media, which together form a pseudostigma. The general lack of
venation except for the three complete plicals is also distinctive. The
simple subcosta is found in most of the groups of the Polpyhagidae.

The tegmina while of normal form, and showing a simple or rela-
tively simple subcosta, have all the other venation incomplete. There
is a short series of anterior radial rami passing to the margin. A
definite posterior apical sector is not present, as this system stops
slightly beyond the mid-point of the tegmina. Both media and cubitus
are fused basally and are reduced to short stubs, the remainder of these
veins being completely lost. Although the plical furrow is present,
no trace of venation may be found here or in the membranous anal field.
The entire apical portion of the tegmina is membranous and wrinkled.

The wing shows one of the most extreme types of specialization
encountered in the Blattaria. The form of the wing is polyphagoid
but a little unusual for the group. The subcosta is short and simple.
The radius separates into Ri and a radial sector; the former soon
thickens and forms a large terminal club occupying an area about
one-sixth the length of the costal margin, while the radial sector ends
abruptly as an incomplete stub. Media rises from the middle of a
definite arculus and soon curves forward and forms a large club, this
club having an area equal to about twice that of the radial club.
Cubitus is present basally and ends abruptly as a bifurcating stub. The
three plical veins are all complete, strong and very prominent. A trace
of a single anal vein is evident. The remainder of the wing is mem-
branous and somewhat wrinkled.

Only the typical genus Holocompsa is placed in this group:

Representative figured: Holocompsa mitidula (Yabricius) figs. 11,
79. [Det. Hebard.] This species is the genotype. Male. Lares,
Puerto Rico.

EUTHYRRHAPHINAE

This subfamily is another of the varied polyphagoid lines of de-
velopment and would appear to be most closely related to the Tiviinae.

JOHN W. H. REHN 41

The tegmina lack practically all venation, so that their relation-
ship cannot be ascertained. However, the tegmina are broader basally
than in most members of the family.

The wings have a shape between that of the Tiviinae and the
Holocompsinae, the plical notch being less evident than is usual for
the family. The subcosta is simple and fused basally with the radius
as in the Tiviinae. zx is present; beyond this there is a well-developed
pseudostigma that obscures the details of the branching of radius.
While this superficially resembles the condition found in the Holo-
compsinae, it is of different origin. Beyond this are a number of
regular apical branches, much as in the Tiviinae. The media has
three well-developed branches, more than in the Tiviinae or other
advanced subfamilies, and is more regular than that in the Polyphagi-
nae. The cubitus has a number of arcuate sectors, the most basal of
which shows secondary branching and twigging as in some of the
Polyphaginae. The plical veins and branched axillary are much as
in the Tiviinae, except that the latter fills over one half of the anal
field instead of about one third.

_ The tegmina are somewhat broader basally than is the norm in this

complex. The anterior margin is weakly arcuate, the apical margin
broadly rounded and the posterior margin weakly arcuate. Traces of
subcosta and the radial stem are the only venation visible in the
densely coriaceous tegima. Both of these veins are represented by rela-
tively short, incomplete, unbranching stems. A sinuate line apparently
marks the plical furrow.

The wing form is slightly unusual for the family in that the plical
notch is scarcely evident. The subcosta is fused basally with radius
and is relatively short, simple and uncurved. The radial system has |
one anterior branch (Ri) and then is obscured by a pseudostigma; be-
yond this four branches are present. The media rises from about the
middle of the well-defined arculus and has three major branches, one of
these being subdivided; all extend to the extreme wing apex. The
cubitus is only moderately simplified having eight primary sectors
which pass diagonally to the wing margins, the most basal showing a
secondary division, and also some twigging towards the plical area.
The first plical vein is unbranched and somewhat sigmoid. The second
plical and third plical veins are entire, strong and direct. Between

MEM. AMER. ENT. soc., 14.

42 CLASSIFICATION OF THE BLATTARIA

these veins, and between them and the associated veins, are single in-
tercalated veins. The first branched axillary is strong and occupies
more than one-half of the anal fan, having five major and some second-
ary branches. Intercalated veins are present in this area. The re-
maining anal veins are simple and incomplete basally.

Only the typical genus is placed in this subfamily.

Representative figured: Euthyrrhapha pacifica (Coquebert) figs.
12, 86. [Det. Hebard, 1920.] This species is the genotype. Male.
Manaos, Brazil.

IDUPILOIMNGARIUDUANIS,

The distinctive genus Diploptera must be placed in a separate
family. The densely coriaceous tegmina which lack distinct venation,
appear superficially to resemble beetle elytra. The very unusual, highly
specialized, wing with its major venation crossing the transverse fold
readily separates this form from any of the other members of the
Blattaria.

The venation of the tegmina, which is for the most part indicated
by clear areas between rows of punctae, is of a simplified type. The
very long subcosta, passing to the apical third is distinctive. The ra-
dius with its simple Ri and anterior rami and lack of an apical pos-
terior sector is comparable to the condition found in some of the higher
Polyphagidae and Blattidae. A single branched media occurs in sev-
eral groups, and the extreme simplification of cubitus, a single stem,
parallels the condition found in the higher Pseudomopinae and their re-
lated forms with appendiculate fields in the wings. The elongate anal
area, set off by a straight plical furrow, is reminiscent of the condition
found in the Panesthiidae.

The wings, with the transverse fold crossing the major venation,
and the division of the radial sector into two simple veins are condi-
tions not found in other Blattaria. The third plical vein supports ap-
proximately one-half of the appendiculate field. In all other forms
with venation in the appendiculate field, such as the Ectobiinae and
Anaplectinae, the third plical is solely responsible for its support.
However, this vein being free basally from the branched axillary is a
condition found in many of the primitive Blattidae. The radius divid-
ing into a distinct Ri and Rs and the second plical vein terminating

JOHN W. H. REHN 43

apically in the first, are conditions found in various groups. The
simple media that is somewhat stalked basally is a feature also found in
the Epilamprinae and other groups. The reduction of cubitus to a
two-branched vein with these branches attempting to reach the apical
margin resembles the condition found in the Pseudomopinae. Thus,
although showing relationship with various subfamilies of the Blat-
tidae, Diploptera presents a combination of features not found in any
one of the stocks.

The tegmina are relatively broad, the anterior margin basally being
strongly convex, then almost straight and roundly passing into the
apical portion. Instead of having the anterio-apical margin the most
produced portion, this arcuately extends to the posterio-lateral angle
which is most produced and is the extreme apex of the tegmina. This
angle is roundly rectangulate. The posterior margin is nearly straight
and passes roundly into the short basal margin. Due to the texture,
very little venation can be seen. The subcosta is heavy and in the basal
sixth angles towards the costal margin, then parallels this margin for a
distance equal to about one-half the total length; no branches or rami
are present. RR: is short, almost parallel to subcosta to the point where
the latter starts to parallel the margin. The radius has at most three
simple anterior rami that go diagonally toward the margin; no apical
posterior branch is present. Media parallels radius to about the mid-
point and then forks into two simple branches, the anterior of which
goes to the apex. The cubitus is a single vein roughly paralleling
media and its posterior branch. The plical furrow is indicated by a
straight line passing diagonally across the posterior portion of the teg-
mina and reaching the margin slightly beyond the end of the subcosta.
Traces of four or five incomplete anal veins are present.

The wings are of very unusual type, and there .is some question
about the interpretation here used. The anterior margin is weakly
arcuate, with a shallow notch at about the middle, this denoting the
transverse fold; the apex is sub-rectangulate, with margins weakly ar-
cuate; posterior margin gradually arcuate to a line behind the costal
notch, which delimits the transverse fold; margin of anal field evenly
arcuate. The subcosta is simple, extending almost to the transverse
fold. Radius with Ri distinct, ending in two or more terminal rami
slightly before the transverse fold. The radial sector is prominent,

MEM. AMER. ENT. Soc., 14.

44 CLASSIFICATION OF THE BLATTARIA

forking before the transverse fold and continuing almost to the ex-
treme apex as two simple branches. The arculus is strong, diagonal.
The media is stalked basally for a short distance on radius: upcurved
slightly before transverse fold, passing over fold and extending almost
to the apex. The post transverse fold portion is rather strongly sig-
moid. The cubitus is strong, almost straight to the transverse fold,
there branching into two simple sectors, one of which extends practi-
cally to the apex, the other ending short of this. The first plical vein
is weakly sigmoid to the transverse fold, passing this fold and ending
slightly before the middle of the appendiculate field. The second plical
vein is relatively strong in its basal one-half and apparently joins the
first plical at the transverse fold. Third plical vein is simple in the
basal half, beyond the transverse fold dividing into two main branches.
The branched axillary has two or three simple branches, remaining ten
anal veins simple.

This wing is folded in the following manner: the anal area folds
fanlike, then the wing folds along the area between the second and
third plical veins. Following this it folds transversely at the middle
with the appendiculate field lying on top of the wing base in repose.

The wing is divided into a considerable number of cells, by the
many cross veins; these are for the most part more or less regular ex-
cept in the posterior half of the appendiculate field, where they are ir-
regular.

This family contains only the nominate genus Diploptera.

Representative figured: Diploptera dytiscoides (Serville) figs. 20,
92. [Det. Hebard.] This is the genotype. Male. Honolulu,
Hawaii.

PANESTHIMIDAE

The Panesthiidae are another distinctive family, but they are more
closely related to the Blattidae than to the other Blattaria. The teg-
mina are long and relatively slender, in texture somewhat resembling
those of the Polyphagidae, but lacking their characteristic venation.
The short, heavy, sigmoid subcosta is reminiscent of the condition
found in the Paranauphoetini and some Pseudomopinae, but in con-
trast to the latter it is equal to one-half or less of the length of the anal
field. The extremely elongate anal field is unusual but is also found in

Sat

JOHN W. H. REHN 4

the Diplopteridae, Oulopterygidae and certain Blattidae. The radial
system, which has anterior and apical rami, is moderately crowded
much as in some of the primitive Blattidae, particularly the Blaberinae.
The retention of a branched media, a primitive feature, is shared with
such forms as the Blattinae. Cubitus is also of a rather primitive type.

The wings are distinctive in that the radial system does not have
more than five branches. This is an unusual condition for the Blat-
taria and a tendency towards it is normally achieved only by extreme
specialization. The lack of a definite arculus in some genera is also
distinctive. On the other hand, the rather long and simple subcosta is
reminiscent of the epilamproids as is also the development of cubitus.
In all forms studied, at least half of the branches of cubitus go to the
plical fold. This is comparable to the condition found in the Epilam-
prinae. The media is usually simple as in many of the Blattidae. In
this small family, the scond plical vein varies from almost complete to
relatively long and terminating in the first plical vein. This is com-
parable to the wide variety of conditions found in various subfamilies
of the Blattidae. The third plical vein normally rises from the
branched axillary, a condition noted in many of the Epilamprinae and
some of the other more specialized groups. The relatively large num-
ber of regular anal veins, in most of the forms, is distinctive. Thus,
the family shows a relationship to many of the larger entities of the
Blattidae.

The tegmina are relatively long and narrow and are usually some-
what membranous; basally they may become somewhat thickened. The
anterior margin is decidedly sinuate. The apical margin is usually
broadly rounded, but at times is subtruncate. The posterior margin is
evenly arcuate, usually with a notch or slight indentation at the plical
furrow. The margin of the anal field is evenly arcuate. The humeral
area is well developed. The subcosta is heavy and short, usually
reaching the margin at the basal fourth. This vein is strongly sinuate
and it is always unbranched. Ventrally it is often produced to form a
tegminal catch. The radius is usually crowded towards the anterior
margin; Ri is not differentiated, and the anterior rami are usually ir-
regular. Near the anterio-apical angle, some rami have secondary
branches. The apical posterior radial branch is well developed, at
times with accessory branches. The media is strong and has two

MEM. AMER. ENT. soc., 14.

46 CLASSIFICATION OF THE BLATTARIA

simple branches. The cubitus has its sectors well developed, but vari-
able; considerable secondary branching is noticed as well as some basal
twigging. In general the branches go toward the posterio-apical angle.
The plical furrow is distinct, strong, lowly arcuate. This furrow
marks off the anal area which is equal to at least twice the length of the
subcosta. The anal field has an indefinite and irregular arrangement
of veins, some showing secondary branching; all are more or less par-
allel to the plical furrow.

The wings have the preaxillary portion rather large, and the anal
area is full. The apical margin is either rounded or subtruncate. A
definite plical notch is present, anal area with its margin broadly and
evenly arcuate. The subcosta is straight and simple, extending for at
least one-half of the wing length. The radial system with Ri distinct
from the sector and extending almost to the apex, always simple.
Radial sector extending for some distance before branching; never
dividing into more than four branches, these may show terminal twig-
ging. The apical posterior branch is very long and simple. All
these branches end at the wing apex. An arculus may or may not be
present. Media, which is usually stalked on the radial sector, is a
simple, direct vein straight to somewhat sigmoid, rarely showing
terminal twigging. The cubitus is strong and has eight or more sec-
tors, at least one-half of these going to the plical fold or fusing apically
and forming an accessory vein parallel to the first plical and the re-
mainder going to the wing apex. The first plical vein is strong and
complete, at times fusing apically with one of the cubital rami. The
second plical vein varies from almost complete and free, to relatively
long and with its apex approximate to the first plical. The third plical
vein is stalked basally on the branched axillary, and its free portion is
complete and simple, reaching the margin behind the plical notch. The
branched axillary normally has two or three well-defined, simple
branches. Behind this there are nine simple anal veins. Intercalated
veins may be present between the major veins, posterior to Ri.

This family has only one apparent line of venational development.
Therefore, the forms studied are all placed in the typical subfamily.

In addition to the typical genus Panesthia, there is at least, the
genus, Salganea.

JOHN W. H. REHN 47

Representatives figured: Panesthia angustipennis (Illiger) figs. 25,
88. [Det. Saussure.] This species is the genotype. Male. Java.
Salganea morio (Burmeister) figs. 19, 89. [Det. Hebard, 1932. |
This species is the genotype. Female. Darjiling District, Himalayas.

BLATTIDAE

The Blattidae are the largest and most varied of the families in-
cluded in the Blattaria. In some respects, primitive venational types
are included within this family as well as a number of highly specialized
types.

The members of this family can be distinguished from most of the
other families by their method of wing folding and support. The anal
area is folded fanwise as in all the families except the Polyphagidae.
When an appendiculate field is present, only in some of the advanced
subfamilies, the field is supported only by the third plical vein; thus the
area is not veinless as in the Oulopterygidae or supported by full vena-
tion as in the Diplopteridae.

The tegmina vary considerably in form and venation, but are never
as densely coriaceous as in the Diplopteridae, nor are they as a rule as
membranous as in the primitive polyphagoids. They usually do not
have as proportionately a long anal area as the Panesthiidae and Dip-
lopteridae. At times, they more or less resemble those of Oulopteryx
in shape and texture, but they never have a subdiamond shaped area
without venation between the radius and media, in the apical one-half,
nor do they normally have a relatively broad area without venation be-
tween the cubitus and the plical furrow. Only the Ceuthobiinae have
the tegmina as membranous as some Polyphagidae and the wing form
will immediately separate these two groups. Those Polyphagidae with
coriaceous tegmina normally have the subcosta much more extensive
than in the present family. The venation varies considerably but
forms with a short subcosta reminiscent of the Panesthiidae, have this
vein equal to more than one-half the length of the anal field. In this
family the anal field is usually either arcuate or angulate apically, in-
stead of being lanceolate. The radial system is crowded anteriorly,
much as in some Polyphagidae, only in some Nyctiborinae and Bla-
berinae. These groups may readily be separated from that family by
a number of features.

MEM. AMER. ENT. Soc., 14.

48 CLASSIFICATION OF THE BLATTARIA

The wings of the Blattidae may immediately be separated from
those of the Polyphagidae by their shape, having an anal area normally
as large as or larger than the preaxillary area. The length of the plical
fold in the Blattidae approximates the wing length, except in some of
those forms with an appendiculate field, while in the Polyphagidae it is
never more than half this length. The Polyphagidae never have an
appendiculate field and the other families that do, Diplopteridae and
Oulopterygidae, either have all major elements of the venation cross-
ing the transverse fold or else lack venation, while in the Blattidae this
field is always supported only by the third plical vein. All the Blat-
tidae have a definite arculus, which is lacking in some of the Panes-
thiidae. The plical veins vary in completeness, but the second one is
rarely complete, and none show terminal twigging, both of which con-
ditions are found in many Polyphagidae. The venation in this family
as a whole is so variable that further comparisons can be made only
within the various entities.

This differential diagnosis should serve to distinguish the Blattidae
from the other families of the Blattaria. They represent a diversified
group for which it is not possible to prepare a concise characterization.
However, the various subsidiary groups may be moderately well de-
fined and their relationships established.

The Blattidae is divided into nine subfamilies which apparently be-
long to four major lines of development. The basic Blattinae and the
Nyctiborinae each appear to represent distinct stocks. The Brachy-
colinae, Blaberinae and Epilamprinae would seem to be derived from
the same basic complex. The remaining subfamilies, Pseudomopinae,
Ectobiinae, Anaplectinae and Ceuthobiinae are also apparently derived
from a common ancestral form.

Tentative Key to the Subfamilies* of the Blattidae
(Alate Forms Only)

1. Wings with apical portion marked off by a transverse fold ............. 2,

Wings without a distinct apical portion marked off by a transverse fold .. 3

2. Tegmina with subcosta simple; wings without an apical posterior radial
branch, apices of major veins joined before the transverse fold.

EcTOBIINAE
Tegmina with subcosta with anterior rami; wings with an apical posterior
radial branch, apices of major veins not joined ....... ANAPLECTINAE

3 The Nocticolinae and Archiblattinae are not included in this key.

JOHN W. H. REHN 49

3. Venation of both tegmina and wings amorphous, the determination of major

VOMS [DSN WNTOSSMTIE ooocaccgccanccoavgdeddoasd0e BRACHYCOLINAE
Venation of both tegmina and wings of more usual type, determination of
MEVOL TSINS Bol WHS IOI MES |DOSSIOO 5 4000000000000 0bnb00guace 4

4. Tegmina with radial system crowded anteriorly, all rami ending at anterior
margin, media either with its branches occupying all or almost all of
the apical margin or they end at anterior margin and cubitus occupies
Eem cipal Mem ay Oni tilaepeayery on ste nen ssiya rate ce sitonsiers aiaeaue avec) Gus eetouss ayalicie es thcsss 5

Tegmina with radial system usually normal, or if crowded anteriorly some
of its branches or rami end at the apical margin, media with its
branches occupying only a portion of the apical margin ............ 6

5. Subcosta of tegmina simple, media ending at anterior margin.

PSEUDOMOPINAE (part)
EUSTEGASTINI
Subcosta of tegmina with anterior rami and a posterior branch, media
occupying most of the apical margin ................. NYCTIBORINAE
6. Third plical vein distinct from the branched axillary and usually remote
ISON IB. 5S:.0 Sere Gre Grats Sete OCI ROR CIC RROIEIE OLS oR IP ean eR Rae hee ene 7

Third plical vein stalked, at least basally, on the branched axillary.
EPILAMPRINAE (part)
7. All or almost all of the cubital branches of the wings going to, or curving
TON ARGMBEMeMTIN AT OTM n rey sie cesy toi ereret succait coat neue tls clave nla temtepmiataad: 8
Numerous cubital branches of the wings going to or ending at the plical
EO LM en eee eyes eae ai aes penta han oy Samet: avertnenel a HiaLgl toe ee A meee sua hw dose 10

8. Cubitus of wings simple, unbranched; margin of area bounding the branched
axillary expanded, practically equal to the preaxillary lobe.

CEUTHOBIINAE
Cubitus of wings with at least one branch; margin of area bounding the
pranchedFaxillanyanoteneathyexpandedi ay aan aaeer create eel 9

9. Wings with media simple, or at most with a single apparently adventitious
branch near the apex; usually all major cubital branches reaching the
margin, rarely most basal branch not reaching this margin (Parco-

VENDA) © S srestau ns thats eo eee Ine Ieee neat rcee PSEUDOMOPINAE (part)
Wings, when not reduced, with the media branched; some cubital branches
MOL Attiamanne WME MAIN ¢oor00cc00c0csgavdo00g4co0ea0nc BLATTINAE

10. Subcosta of tegmina always considerably longer than anal area, always with
anterior rami and a posterior branch; wings with accessory veins in
the base of the plical area, radius appearing to be crowded toward the
DALE IL OxApI Cale SSG my Meer eter eta etat funteteya tec asiele sere alien BLABERINAE

Subcosta of tegmina shorter than, to slightly longer than, anal area, with
or without anterior rami and but rarely with a posterior branch; wings
without accessory veins in the base of the plical area, radius not appear-
ing to be crowded toward the anterio-apical angle ................ 11

MEM. AMER. ENT. Soc., 14.

50 CLASSIFICATION OF THE BLATTARIA

11. Subcosta of tegmina with a few short anterior rami apically, no posterior
apical radial branch, media four branched; wings with a small inter-

plical field, second plical vein ending free .... PSEUDOMOPINAE (part)

ISCHNOPTERINI

Without this combination of features ............. EPILAMPRINAE (part)
SP UA RINNE CEUTHOBIINAE

BRACHYCOL INAE BLABERINAE ANAPLECTINAE

ECTOBIINAE

NYCTIBORINAE

PSEUDOMOP INAE
LATTINAE

ANCESTRAL STOCK

Diagram 3.—Probable relationship of the components of the Blattidae.

As shown in the diagram, the Blattinae are considered to be the
most primitive of the existing Blattidae with three major lines of de-
velopment. The two which are composite have been termed, for the
sake of convenience, the Epilamproid complex and Pseudomopoid
complex.

BLATTINAE

The Blattinae are the most primitive of the components of the
Blattidae. The tegmina usually have a moderately developed humeral
area as in the pseudomopoids and some of the epilamproids. The sub-
costa is rather heavy and may have anterior rami as in the epilamp-
roids and certain pseudomopoids, but never has the posterior branches
found in some of the epilamproids and the Nyctiborinae. The radius
has irregular anterior and apical rami much like those in the epilampro-
ids and not as regular as those in the pseudomopoids. The media is

JOHN W. H. REHN Sil

branched, always free from cubitus as in the epilamproids and the
primitive pseudomopoids and not enormously developed as in the
Nyctiborinae. The cubitus has some indication of two stems, its
branches varying from oblique to tending to parallel the posterior mar-
gin, thus showing affinities with the pseudomopoids on one hand and
the epilamproids on the other. The plical furrow is evenly arcuate and
the anal veins are more or less transverse with some branching apically
and posteriorly as in the epilamproids.

The wings never have an appendiculate field as in the Anaplectinae
and Ectobiinae. The wings have the subcosta simple or with a termi-
nal fork, relatively short as in the pseudomopoids. Radius is well de-
veloped with some anterior rami, but these are less regular than in
the pseudomopoids and are more numerous than in the epilamproids.
The branched media is much as in most epilamproids, not simple as in
the pseudomopoids and Nyctiborinae. The second plical vein is usually
relatively long and free apically thus like some epilamproids and some
pseudomopoids, but never fused apically with the first plical as often
happens in both of these groups. Usually, it is not as long as in the
Nyctiborinae. The third plical vein is free and remote from the
branched axillary as in the pseudomopoids and the Nyctiborinae. The
branched axillary is relatively simple and remote from the anal veins
as in both the pseudomopoids and the Nyctiborinae, thus it is markedly
distinct from the anterior fan.

The tegmina, in general, are elongated oval except when reduced,
either by truncation or to a lobiform flap, and even then retain most of
the essential venation, but on a reduced scale. In texture they vary
from membranous, transparent to somewhat coriaceous. The over-
lap is but weakly indicated and then only in the more highly evolved
genera. The anterior margin is either practically straight or weakly
convex, roundly passing into the usually arcuate apical margin. In
forms showing reduction, the anterio-apical angle is sub-rectangulate.
Apical margin roundly passing into the convex posterior margin. The
plical furrow may or may not be indicated by a shallow notch. Anal
field margin arcuate and passing by an obtuse angulation into the basal
margin. The humeral area is well developed, except in Pseudodero-
peltis. The subcosta is usually heavy, either moderately sinuate or
nearly straight, reaching the anterior margin at or near its greatest

MEM. AMER. ENT. Soc., 14.

a2 CLASSIFICATION OF THE BLATTARIA

anterior expanse, and about as far apically as the end of the plical fur-
row. It is reduced in genera showing tegminal reduction. Some ir-
regular rami are present in the more primitive genera, but they are
absent in the more evolved forms. The radius is moderately well de-
veloped and has a number of anterior rami, probably including the un-
differentiated Ri, some simple, others with more or less irregular
secondary branches. Apically these branches tend to be dichotomous.
The radial sector is usually well developed, with one or more branch-
ing anterior rami; an apical posterior sector is present, but often it is
not well differentiated. The media is free basally and well developed,
having four or more major branches, and with terminal subdivisions,
except in those forms showing considerable reduction; often with ad-
ditional incomplete basal branches. In some specialized genera media
curves away from radius. The cubitus is well developed and usually
has two main stems each with numerous supplementary divisions
(branches). There are usually at least eight of these branches which
in general go diagonally toward the apico-posterior angle. The plical
furrow is distinct, usually gradually and variably curved to posterior
margin. A plical notch may or may not be present. The anal field
has a considerable number of veins which more or less parallel the
posterior margin, some of the more posterior ones branched. Inter-
calated veins are present in the vicinity of the radial sector, and in the
median and cubital fields, except in genera showing considerable re-
duction.

The wings have a relatively large preaxillary area and a well-de-
veloped, but considerably shorter anal area. The anterior margin
varies from straight to moderately convex and the apical margin is
broadly and relatively evenly rounded, except in those forms showing
reduction. A definite notch is present at the plical fold even in genera
showing reduction. The margin of the anal field is arcuate, roundly
passing into the basal margin. The subcosta is simple, straight to
slightly curved and extends from one-third to one-half the wing length,
except in the genus Pseudoderopeltis where it is shorter. Ri primi-
tively has four or more rami, but a reduction occurs and some genera
have this vein unbranched. Radius has relatively numerous anterior
and apical rami, the more apical ones often subdividing. An apical
posterior sector is present and often has a few branches. In forms

JOHN W. H. REHN 53

having reduction, it may not be distinctly differentiated. Media is
primitively branched and free, rising from the apparent top of the arcu-
lus, but in the more advanced forms becomes stalked on the radius.
The details of branching vary, but all genera not having marked re-
duction have at least two major branches and these usually subdivide.
The arculus is strong, relatively prominent, and usually oblique. The
cubitus usually has about seven main branches with numerous second-
ary divisions which go towards the apical margin. There is a marked
tendency for these branches to fuse apically, in some cases basally; or
median anastomosis may occur. The area below the arculus is free of
basal twigging. The first plical vein is free, simple and complete;
apically it is either straight or curved slightly forward. The second
plical vein is free, simple and, from long and almost complete, to mod-
erately short, but the apex is always free. The third plical vein is
strong and complete; its origin is free from the branched axillary. The
branched axillary is strong, its base free from the third plical, and dis-
tinct from that of the other veins and it has from four to six branches,
some showing a secondary branching. The anal fan is well developed
with eight to twelve veins, exclusive of the branched axillary. Rarely
some of these may show secondary branching.

Intercalated veins are present in the median field, and throughout
most of the cubital field, rarely, as in Deropeltis, they are present in
the radial field. The male of Blatta having the wings reduced lacks
these.

An examination of a limited number of genera of this subfamily
shows a diversity in development. A more comprehensive survey will
be necessary to determine whether or not tribes may be represented.
Both Periplaneta and Deropeltis have a variety of primitive features,
and it seems probable that Blatta and possibly Ewrycotis have evolved
from this same ancestral stock. On the other hand, Pseudoderopeltis
in some respects appears moderately distinct.

Representatives figured: Blatta orientalis Linnaeus figs. 24, 29, 93.
[Det. Hebard.| This species is the genotype. Male and Female.
Bloomington, Indiana.

Deropeltis erythrocephala (Fabricius) figs. 26,96. [Det. J. A. G.
Rehn, 1920.] This species is the genotype. Male. Transvaal.

MEM. AMER. ENT. soc.. 14.

cn
ass

CLASSIFICATION OF THE BLATTARIA

Eurycotis floridana (Walker) fig. 28. [Det. Hebard.] The
genotype is Polyzosteria rufovittata Brunner [= Eurycotis mysteca
(Saussure) ]. Male. Biscayne Bay, Florida.

Periplaneta americana (Linnaeus) figs. 21, 94. [Det. Hebard.]
This is the genotype. Female. Trichinopoly, Madras, India.

Pseudoderopeltis brevicollis (Serville) figs. 27,95. [Det. J. A. G.
Rehn, 1920.] The genotype is Deropeltis antennata Saussure
[—= Pseudoderopeltis antennata (Saussure) |. Male. Pretoria, Trans-
vaal.

NYCTIBORINAE

The Nyctiborinae represent a distinct group of the family and ap-
pear to be related to all its other major components. The tegmina are
relatively broad, richly veined and have a small anal area. They have
a large humeral area, and the subcosta, with anterior rami and pos-
terior branches is as in some of the epilamproids. The radial system
has the rami more crowded and the whole system terminates at the
anterior margin, a distinctive feature, as in all the other groups at least
some part of this system attains the apical margin. In conjunction
with this the media has been greatly developed, more so than in any
other existing group, its numerous branches occupying all or almost all
of the apical margin. The cubitus is inclined to divide into two stems
and the sectors may be oblique and go toward the posterio-apical angle
or tend to parallel the posterior margin, thus agreeing in part with the
Blattinae and also with the pseudomopoids. The plical furrow is
evenly arcuate as in some of the Blattinae and in epilamproids, but
marks off an anal area proportionately smaller than that usually found
in any of the other groups. The veins in this area tend to follow the
curve of the nearest margin, with those mesad more or less straight as
in the epilamproids, or there may be some anterior branching reminis-
cent of the condition found in certain of the Polyphagidae.

The wings, which are richly veined, have a large preaxillary area
and a shorter anal area, much as in the Blattinae. The subcosta is
either long and almost simple or with a few rami, resembling both the
Blattinae and epilamproids. The radius is well developed, with nu-
merous anterior and apical rami and has a well-developed apical pos-
terior branch thus resembling, but richer than, the Blattinae. The

JOHN W. H. REHN

on
un

media is simple as in the pseudomopoids and many epilamproids. Cu-
bitus has most of its sectors going to the apex and the remainder to the
plical fold, much as in the primitive epilamproids. The second plical
vein varies from complete to long and free apically, in the former re-
sembling the Polyphagidae and certain Blaberinae while in the latter it
is but little longer than in the Blattinae. The third plical vein is remote
from the branched axillary which is distinctly separated from the re-
maining anals, resembling the condition found in both the Blattinae
and the pseudomopoids.

The tegmina are broadly ovate with the anterior margin convex,
passing into the broadly rounded apical margin and roundly passing
into the subtruncate posterior margin. The plical notch is not well
developed. Margin of anal field at first weakly arcuate or subtruncate,
angulately passing into the basal portion. The humeral area is
broadly expanded. The subcosta is heavy and either straight or angu-
late, approximating in length the anal field, with numerous to abundant
anterior rami, some of which secondarily branch. Posterior rami,
representing captures from the radial system, prominent, subdivided.
The radius has numerous to extremely numerous irregular anterior
rami, many with secondary and further branching; apical rami sub-
dividing even further. No well-differentiated apical posterior sector
is present. All radial branches go to the anterior margin of tegmina.
The media with its origin near radius, soon divides into a large number
of sectors with numerous secondary and further branches. Posteriorly,
it is practically impossible to differentiate media from cubitus, these
branches filling all or nearly all of the apical margin. The base of
cubitus is close to that of media; there are usually two major stems
each with a number of branches which further subdivide. These
numerous sectors are either oblique or tend to parallel the posterior
margin. The plical furrow is strongly arcuate and the plical notch is
poorly developed. The small anal area has numerous veins, some of
which show branching.

Wings with the preaxillary area large and anal area relatively
small. The subcosta is equal to one-half or less the wing length,
usually with irregular branches. The radius with a distinct Ri, which
after paralleling subcosta, has anterior rami. The remainder of the

MEM. AMER. ENT. soc., 14.

56 CLASSIFICATION OF THE BLATTARIA

radial system has numerous irregular, often secondarily branched an-
terior and apical rami. An apical posterior branch with secondary
divisions is present. The media, which is stalked for a short distance
on the radial sector, is a simple vein, with or without terminal twig-
ging, straight or arcuate. The arculus is strong and oblique. The
cubitus has a number of irregular sectors of which one-half or more,
but usually two-thirds, go to the apical margin and the others to the
plical fold. Many of these branches have irregular secondary divi-
sions. Some basal twigging is present near the arculus. The first
plical vein is free, distinct and straight. The second plical vein is free,
distinct, either reaching wing apex or nearly so. The third plical vein
is free, distinct, straight, passing to the margin at or behind the plical
notch. The branched axillary is strong, free from, and curving away
from, the third plical; usually, it has four major branches, and some
irregular secondary branches. A large number of simple anal veins
are posterior to, and free from, the axillary.

The members of this subfamily are readily divisible into two com-
ponents, those related to the typical genus, Nyctibora and the unusual
Megaloblatta. It is believed that those entities are sufficiently dis-
tinct to be considered tribes. The Nyctiborini have a much less
branched venation than is found in the Megaloblattini. The former
have the tegmina with the subcosta straight, and with only a moderate
number of rami while in the latter, the subcosta is strongly angled and
has abundant rami. In the Nyctiborini, the radius, media and cubitus
have fewer branches and less subdividing than in the other tribe.
Also, in the Megaloblattini the sectors of cubitus are oblique instead of
paralleling the posterior margin. The wings present the same general
features, fewer branches, and branching in the Nyctiborini. Inter-
calated veins are found only in the Megaloblattini.

Nyctiborini

The members of this complex may be separated from the Megalo-
blattini by the features already mentioned. In addition the tegmina
have the humeral area more produced basally and its margin is defi-
nitely angulate instead of arcuate. The subcostal rami are few and
terminal, not widespread. The posterior branches of subcosta are
fewer and have only single dichotomous branching. There is much

7

or

JOHN W. H. REHN

less secondary division of the radial rami and the radius is more re-
stricted in its area. Both media and cubitus are more regular in their
branching and these attempt to parallel the posterior margin. The
anal field shows very little branching and the veins tend to parallel the
nearest margin.

The wing in this tribe has the anal area proportionately larger than
in the Megaloblattini. The subcosta is more regular and the radius
has fewer branches. The media is straight instead of arcuate and
cubitus has considerably fewer and more regular sectors. The second
plical vein is not complete. The branched axillary is not as remote
form the third plical vein and does not show any secondary divisions.
There are more simple anal veins than in the other tribe.

It does not appear to be necessary to further characterize this tribe,
to which most of the members of the Nyctiborinae belong.

Representative figured: Nyctibora noctivaga Rehn figs. 22, 90.
[Det. J. A. G. Rehn, 1920.] The genotype is Nyctibora sericea Bur-
meister. Male. Changuinola District, Panama.

Megaloblattini

This tribe which is formed for the nominate genus has the follow-
ing distinctive features, in addition to those previously noted. The
tegmina have the humeral area less expanded, and its margin is arcu-
ate instead of angulate. The subcosta, which is angulate, has a full
series of rami, some of which have secondary branches and the pos-
terior branches are more numerous and less regular than in related
forms. The radius, media and cubitus all are exceedingly branched
and all of these have numerous subdivisions. The branches of cubitus
are oblique instead of parallel to the posterior margin. The anal field
has the veins in its anterior portion with a number of branches, much
as in some Polyphagidae. The remainder of the veins are more or less
oblique and show some branching. The anal field is proportionately
smaller than in the Nyctiborini.

The wing, which has a complete series of intercalated veins, has
the anal field proportionately smaller than in the other tribe. The
radius is more irregular and has more numerous branches. Media is
arcuate instead of straight. The cubitus shows.more branching both
primary and secondary and is more irregular. The second plical vein

MEM. AMER. ENT. Soc., 14.

58 CLASSIFICATION OF THE BLATTARIA

is complete. The branched axillary is free, and relatively remote from
the third plical vein; its branches have some secondary divisions.
Only the primitive genus Megaloblatta is placed in this tribe.
Representative figured: Megaloblatta blaberoides (Walker) figs.
23, 91. [Det. Hebard, 1919.] The genotype is Megaloblatta peru-
viana Dohrn. Male. Muzo, Boyaca, Colombia.

EPILAMPROID COMPLEX

The epilamproid complex includes three subfamilies, Brachycol-
inae, Blaberinae, Epilamprinae, all of which appear to have evolved
from the Blattinae. Many have the humeral area of the tegmina de-
veloped as in the Nyctiborinae, while some retain this area much as in
the Blattinae and in pseudomopoids. The subcosta, while primitively
like that of the Blattinae, more often has numerous anterior rami and
some members have acquired posterior branches as in the Nyctibor-
inae. The radius may be of the type found in the Blattinae, but often
with irregular and oblique rami, and with terminal branches. The
apical posterior branch is often well developed and with more branches
than in the other groups. Primitively, there is a well-branched media
as in the Blattinae, but in advanced forms it is reduced; rarely fused
with cubitus, as in some members of the pseudomopoid complex, or
tremendously developed as in the Nyctiborinae. Cubitus primitively
with numerous branches, at times with two stems as in many Blattinae,
the sectors mostly go toward the posterio-apical angle. Advanced
forms have fewer branches, but they rarely parallel the posterior mar-
gin as in many of the pseudomopoid complex. The plical furrow is
evenly arcuate as in the Blattinae, not limiting a small anal area as in
the Nyctiborinae or angulate as in many of the pseudomopoid complex.
Anal veins normally tend, more than in the other groups, to parallel
the furrow. They may be branched as in the Blattinae.

Wings with the subcosta longer than in the other groups, and often
simple. Ri well differentiated, usually with more rami than in the
pseudomopoids, much as in the Blattinae and Nyctiborinae. Radial
sector with some anterior and apical rami, the apical posterior branch
is often well developed, primitively as in the Blattinae, but the advanced
forms show reduction. Media branched in Blaberinae, Brachycolinae
and in some Epilamprinae, in others simple as is also the case in the

JOHN W. H. REHN 59

Perisphaerini. The former condition is comparable to that in the
Blattinae, while the latter is like that in the pseudomopoid complex and
the Nyctiborinae. The cubitus is primitively as in the Blattinae and
Nyctiborinae with a few branches to the plical fold, specializing by
having more branches to this fold until in the Thoracini all but the last
two do not reach the wing apex. The first plical vein is usually straight
as in the Blattinae, rarely curved forward as in the pseudomopoid com-
plex. In some, it is fused with the coalesced tips of the cubital sectors,
a condition practically distinctive of this group. The second plical
vein varies from long to short and its apex may be free or not. In
this some genera resemble the Blattinae and others some of the pseudo-
mopoid complex, but in none is it as long as in the Nyctiborinae. The
third plical vein is either free, or fused for a varying distance with the
branched axillary, never free and remote from this vein as in the Blat-
tinae, Nyctiborinae and the pseudomopoid complex. The branched
axillary is either comparable in size to that of the Blattinae and of the
pseudomopoid complex, or it may be more developed and support up
to one-half of the anal field.

EPILAMPRINAE

The Epilamprinae is the largest and most varied in the complex
and as here understood includes not only the Epilamprinae of authors,
but also the Panchlorinae, and Perisphaerinae and certain other con-
cepts which have been put in other subfamilies. This assemblage in-
cludes a number of diverse lines of specialization, but apparently is of
homogenous origin. The diversity is such that a number of tribes
must be recognized.

The tegmina may have the humeral area but moderately developed
as in the Blattinae or it may be large, as in the other members of this
complex. The development of the rami of subcosta indicates a similar
relationship; posterior branches are also present at times, as in the
Blaberinae. The radius has more branches and is more irregular than
in the Blattinae, except in the Perisphaerini, somewhat as in the
Blaberinae, although the field is not as broad as in that subfamily. The
radius is not as irregular as in the Brachycolinae. Media is mod-
erately branched as in the Blattinae, particularly in the Perisphaerini.
It has fewer branches than in the Blaberinae, and only rarely is simple

MEM. AMER. ENT. soc., 14.

60 CLASSIFICATION OF THE BLATTARIA

as in some Pseudomopinae. The cubitus often has two main stems
with secondary branches. These may be regular or irregular, and go
obliquely to the posterio-apical angle, agreeing in this respect with the
Blaberinae and Brachycolinae. The more anterior anal veins tend to
parallel the plical furrow, while the posterior ones are more or less
transverse, resembling the condition found in the Blattinae.

Wings with the subcosta long and simple as in the other subfam-
ilies. Ri about as in the Blattinae but longer, usually not as long as
in the Blaberinae. The media is usually simple as in many other
groups. Cubitus with at least one-half of its branches going to the
plical fold, often more; this is practically distinctive. The second plical
vein usually terminates in the first plical, not free as in the Blaberinae
and Brachycolinae, but it is short in the Perisphaerini. The third
plical vein is free or fused with the branched axillary, never remote as
in the other groups. All members of the group with anterior branches
from the branched axillary, and this vein is not remote from the re-
maining anal veins.

The tegmina vary from relatively short and broad to rather long
and slender, however, they never approach the extreme condition found
in certain Blattinae and Panesthiidae. The anterior margin is gen-
erally weakly convex, but varies from almost straight to sigmoid.
The apical margin is usually broadly rounded, at times only narrowly
so, rarely almost subrectangulate. The posterior margin varies from

weakly arcuate to almost straight and a notch is usually present at the
“apex of the plical furrow. The margin of the anal field is occasionally
subangulate basally. The humeral area is usually at least moderately
well developed, in some genera it is greatly expanded; rarely it is re-
duced. This latter condition is found in some of the more specialized
genera. The subcosta is strong, usually approximating in length the
anal field, always with some anterior branches. Basally it is heavy
and thickened, at times this tendency is noted in some of the anterior
rami. Anterior rami normally rather irregular and numerous; rarely
showing secondary banches, at times reduced. In at least some of the
subdivisions posterior rami are present. Radius with at least a mod-
erate number, often abundant, anterior rami; at times some of these
stalked on the subcosta. These vary from very irregular and with
secondary branches to regular, spaced and simple. In general, a tend-
ency toward reduction and simplification of radius is noted in the more

JOHN W. H. REHN 61

advanced forms. Usually some apical rami, which may have secondary
divisions, are present. An apical posterior radial branch is usually
present although it is often poorly developed or not well differentiated.
Rarely, the radius appears to be stalked with a portion of media. The
media primitively has four regular branches which extend to the apex.
However, there is a gradual reduction in branches as well as an ap-
proximation of the basal portion of media to the radius. In certain of
the complexes, the base of media has become involved with that of
cubitus so that they can not be separated. The cubitus primitively is
rather irregular with numerous branches going diagonally to the pos-
terio-apical margin. The division of this vein into two stems is often
evident. In more advanced members of the group certain tendencies
are noted, one the addition and another the reduction in the number of
cubital branches. In both of these cases branching is regular with
little secondary division, and the branches tend to more or less parallel
the posterior margin. When these conditions are found, the division
into two main stems is usually not evident. The plical furrow is dis-
tinct, with at least a noticeable apical notch. This furrow is strongly
to weakly arcuate, never distinctly angulate; usually, it more or less
parallels the basal cubital branch. The anal area is broad and about
as long as the subcosta. It has a moderate to large number of simple
veins, which in general go arcuately across the field.

The wings show considerably more diversity than do the tegmina,
although certain basic similarities exist among all the members of this
subfamily. Certain distinctive venational features are taken as in-
dicating tribal division. The anterior margin is usually evenly arcu-
ate, rarely subsigmoid and the apical margin varies from broadly to
rather narrowly rounded, rarely subtruncate. The plical notch is nor-
mally evident, at times strong, rarely subobsolete. Margin of anal field
usually evenly and broadly arcuate, at times angulate basally. The
anal field is large. The subcosta is always simple and more or less
parallels the costal margin, usually ending past the midpoint, at times
extending almost to the apical third. Radius with Ri always well
differentiated and having anterior rami; these are irregular in some
groups, but, usually, rather regular and rarely showing any secondary
division. The remainder of radius parallels Ri, then has some an-
terior and apical rami; these are regular or irregular. Often there is

MEM. AMER. ENT. soc., 14.

62 CLASSIFICATION OF THE BLATTARIA

a short, simple apical posterior radial branch going to the apical margin,
rarely directed obliquely posterior. The arculus is always strong, and
usually oblique. The media normally apparently rises from the top or
the middle of arculus, rarely stalked on radius. Usually it consists of
a simple arcuate or sigmoid vein, at times having some terminal branch-
ing. The cubitus is strong and has a large number of oblique
branches; usually more than half of these go to the plical area, while
the remainder go to the apex. Although these are at times rather ir-
regular, extreme regularity is found in most of the members of the
group. The arrangement of these sectors is a distinctive feature of the
group. The first plical vein is always simple and complete, at times
slightly upcurved apically, not joining any of the cubital branches.
The second plical vein varies from long and free apically to, in the
majority of cases, short and terminating in the first plical. The third
plical vein is often stalked upon the branched axillary and except in ex-
treme modifications is straight and goes to the margin behind the plical
notch. Usually at least ten veins are present in the anal area, vary-
ing from eight to more than sixteen. The branched axillary, which is
sometimes fused basally with the third plical, normally has from two to
five simple branches. It is not always possible to tell which is the most
posterior branch of this vein because of the arrangement of other anals
along the basal bar. Intercalated veins are always present in the anal
area and may appear in the preaxillary portion.

This subfamily may be divided into the following tribes : Calolamp-
rini, Epilamprini, Leucophaeini, Litopeltini, Nauphoetini, Oniscosom-
ini, Panchlorini, Paranauphoetini, Paratropini, Perisphaerini, Phoras-
pidini and Thoracint.

Calolamprini

This small tribe has a combination of features that serve to sepa-
rate it from related forms. As the organs studied are not fully de-
veloped in the female sex, the discussion concerns only males. The
tegmina resemble in most respects those of the Oniscosomini. The
simpler, heavier, and shorter subcosta which is considerably shorter
than the anal area, will serve to distinguish this tribe. The branches
of cubitus are more irregular than in any other tribe.

’

JOHN W. H. REHN 63

NAUPHOET INI

EPILAMPRINI
PANCHLORINI

L JO OPIEILY JINN

THORAC INI
CALOLAMPRINI

PARANAUPHOET INI
PERTSPHAERINI

ONISCOSOMINI PHORASPIDINI

PARATROPINI
LEUCQPHAE INI

ANCESTRAL STOCK

Diagram 4.—Probable relationship of the components of the Epilamprinae.

The wings are more like those of the Panchlorini, having the anal
area more produced than in the Oniscosomini and Leucophaeini. The
subcosta is shorter than in most of the other tribes. Ri has spaced
anterior rami, and the cubitus has only about one-third of its branches
going to the apex, as in many of the more highly evolved groups.

The tegmina are relatively broad with the apex evenly and broadly
rounded, more so than in most of the other tribes. The subcosta
which is equal to only about two-thirds the length of the anal field is
very heavy and has only four spaced, simple anterior rami. The
radius curves forward slightly before the middle. It has, basally,

MEM. AMER. ENT. Soc., 14.

64 CLASSIFICATION OF THE BLATTARIA

regular anterior rami, then ones with secondary branching. Irregu-
lar apical branched rami are also present as is a short, simple, apical
posterior branch. The media parallels radius and has at least three
simple branches. ‘These start near the middle of the tegmina, approxi-
mate to the forward curve of radius. The cubitus has several irregu-
lar, oblique branches. The plical furrow is arcuate, and a shallow
plical notch is present. The anal field has a few veins, the more me-
dian of which branch.

The wings have a proportionately smaller preaxillary area than in
the Oniscosomini and Leucophaeini. The anal field is fuller and more
produced, projecting almost as far as the preaxillary portion, more as
in the Phoraspidini and Homalopteryx. The subcosta is simple and
shorter than in most other groups, reaching the anterior margin before
the middle. R: has only four regular anterior rami, much as in the
Leucophaeini. The radius has a few apical rami and the apical pos-
terior branch is not differentiated, which is unusual. The media is
simple and arcuate. The cubitus has moderately regular oblique
branches of which one-third go to the apical margin. The first plical
vein is simple and direct, not upturned apically. The second plical vein
is very short and ends in the first plical. The third plical vein is free
from the branched axillary, simple and direct. No interplical field is
present. The branched axillary has three simple branches and there
are nine simple veins posterior to it. Intercalated veins are found in
both the preaxillary area and the anal field.

At present only the typical genus is placed in this tribe.

Representative figured: Calolampra irrorata (Fabricius) figs. 30,
97. [Det. Saussure.] This species is the genotype. Male. “ Nou-
veau Holland” [= Australia].

Oniscosomini

This distinctive tribe is apparently most closely related to the
Leucophaeini and Calolamprini. As females of this tribe have the teg-
mina and wings markedly reduced the discussion is based on the con-
dition found in the males. The amorphous venation of the tegmina is
comparable only with that of the Calolamprini. The subcosta with-
out a complete series of anterior rami, the freely branched media and
the oblique cubital branches are all primitive features shared with some

on

JOHN W. H. REHN 6

of the other members of the subfamily. The anal field approximates in
length the subcosta, which again is a primitive condition.

The wings have the anal area considerably shorter than the pre-
axillary area, as in the Leucophaeini. The irregular venation is dis-
tinctive of this tribe. In general, there are fewer cubital branches than
in the other tribes. The first plical vein is straight apically and no
plical field is present, agreeing in these respects with some of the other
tribes of the group. The branched axillary with the most anterior
branch rising first is distinctive.

The tegmina vary from relatively broad to rather slender,
rounded apically much as in the Leucophaeini, but less so than in most
of the other members of the subfamily. The humeral area is moder-
ately or distinctly produced. The subcosta is heavy basally as in the
Epilamprini and Leucophaeini, and has a small number of irregular,
and at times secondarily branched, anterior rami. These are absent
basally and the most apical ones may be connected with some of the
anterior radial rami. The radius has a number of irregular, second-
arily dividing anterior rami and the irregular apical rami also have
subdivisions. A distinct, simple, posterior branch is present, usually
rising slightly past the middle. The media is more remote from the
radius basally than in most members of the group, and it divides into
three or more distinct branches, some of which have terminal sub-
divisions. The cubitus has a definite anterior branch as in the Pan-
chlorini and some Epilamprini. The remainder divides into a number
of regular sectors, some dividing secondarily; these go obliquely to-
ward the postero-apical angle. The plical furrow is distinct and an
apical notch is at least indicated. Although this furrow is close to the
basal portion of cubitus the latter does not curve evenly around it as in
certain of the other tribes, agreeing in this respect with the Leucophae-
ini. The anal area has a number of irregular veins the more median
of which usually branch, as in several other tribes.

The wings have a large preaxillary area, a well-defined plical notch
and an anal area proportionately smaller than in most of the other
tribes. The subcosta is long, extending to almost the apical third,
simple and direct. The radius divides into a distinct Ri which has
some irregular terminal twigging, much as in some Leucophaeini.
The remainder of radius is practically simple to below the apex of the

MEM. AMER. ENT. Soc., 14.

66 CLASSIFICATION OF THE BLATTARIA

subcosta then it has some irregular apical rami. In the genus Blaptica,
a definite posterior radial branch is present, while this is greatly re-
duced in Oniscosoma. The radius is often connected with Ri by some
adventitious crossveins. The arculus is strong and diagonal. Media
is either sinuate or arcuate and goes to the extreme apex; in Omts-
cosoma, it has simple terminal branches. The cubitus is strong and
has a number of sectors, usually seven to ten, of which about one-half
go to the apical margin, the remainder towards the plical furrow. At
times their apices curve and join. The first plical vein is free and
straight. The second plical vein is relatively short, equal to between
one-third and one-half the wing length, and always ends in the first
plical. The third plical vein may be stalked, at least for a short dis-
tance, on the branched axillary, and goes to the margin slightly behind
the plical notch. The interplical area is not expanding. The branched
axillary normally has three branches, the more anterior one rising first,
which is distinctive of this tribe. Eight to ten simple anal veins are
posterior to the branched axillary.

It is believed that a number of genera in addition to those studied
belong to this tribe.

Representatives figured: Blaptica dubia (Serville) figs. 32, 99.
[Det. Hebard, 1921.] The genotype is Blaptica claraziana Saussure
= Blabera dubia Serville [==Blaptica dubia (Serville)]. Male.
Buenos Aires, Argentina.

Oniscosoma granicollis (Saussure) figs. 31, 98. [Det. J. A. G.
Rehn, 1922]. The genotype is Oniscosoma castanea Brunner = Ze-
tobora granicollis Saussure = [Oniscosoma granicollis (Saussure) ].
Male. Dunkeld, Victoria, Australia.

Perisphaerini

On the basis of wing venation, this group, which is usually con-
sidered to be at least a distinct subfamily, can only be regarded as a
tribe of the highly developed Epilamprinae. The group appears to be
most closely related to the Oniscosomini and Calolamprini.

The tegmina are distinctive with their strongly angulate plical fur-
row, a condition not found elsewhere in the Epilamprinae, but which
occurs in the Pseudomopinae. The large humeral area which lacks
venation in its basal portion is comparable to that of the Oniscosomini

JOHN W. H. REHN 67

and some other tribes. Posterior subcostal branches are absent as in
most of the lower Epilamprinae. The arrangement of the regular an-
terior radial rami resembles the condition found in the more advanced
tribes. A simple, long, apical posterior branch is present as in the
Oniscosomini. The separate origin and then fusion for a short dis-
tance of cubitus and media, is distinctive.

The wings resemble those of the Oniscosomini in many respects,
but Ri is shorter than in that group. The second plical vein is short
and ends in the first plical, much as in the Calolamprini. The third
plical vein is stalked on the branched axillary as in many of the tribes,
and this latter vein has only one definite branch as in many of the
Epilamprini and Litopeltiini.

The tegmina are rather broad and short, almost subrectangulate.
The anterior margin is weakly arcuate and rounds into the broadly
arcuate apical margin. The posterior margin is weakly concave and
the plical notch is not evident. The margin of the anal field has a
rounded angulation basally. The humeral area is well developed.
The subcosta is heavy and oblique, with a few spaced rami in its apical
half. The radius has a number of anterior rami which rarely have
secondary divisions. The apical rami are not well developed. A

g, simple apical posterior branch is present. The media soon be-
comes fused with cubitus, then free, and has apparently at least four
branches some of which may show secondary subdivisions. The cubi-
tus divides into two main stems each of which have numerous sub-
divisions. These branches are arcuate or sigmoid and in general go to
the postero-apical margin. It is not possible to completely determine
the limits of these two systems. The plical furrow is strongly arcuate
and delimits a large anal area. Basally the furrow is evenly arcuate,
then it has an obtuse angulation; from this to the posterior margin it
is arcuate. The subrectangulate anal area has a few simple, relatively

long

straight veins.

The wings have a relatively large preaxillary area and a smaller,
moderately developed anal fan. The anterior margin is sigmoid and
the apical margin is rounded. A shallow plical notch is evident and
the anal field margin is evenly arcuate. The simple subcosta is long,
reaching the margin beyond the middle. The radius has a distinct Ri
which basally parallels the subcosta, beyond that it has a few apical

MEM. AMER. ENT. soc., 14.

68 CLASSIFICATION OF THE BLATTARIA

rami. The radial sector is strong and has at least traces of some apical
rami. No distinct posterior apical branch is present. The media is a
simple vein which is stalked basally on the radial sector. The arculus
is transverse. The cubitus is well developed with a number of oblique
more or less arcuate sectors, more than half of which go to the apical
margin. There is some secondary division of these branches. The
first plical vein is strong and straight. The second plical vein is rela-
tively short and ends in the first plical vein. The third plical vein is
apparently stalked for a short distance on the branched axillary, then
becomes free and goes directly to the margin. No interplical field is
present. The branched axillary is simple having only one well-defined
branch and behind this there are nine simple anal veins. Throughout
the anal field there are short intercalated veins.

Only the typical genus has been examined, but it is believed that a
number of other genera can be included in this complex.

Representative figured: Perisphaerus contiguus (Burmeister) figs.
33, 100. [Det. Hebard, 1932.] The genotype is Perisphaerus arma-
dillo Serville. Male. Mount Lamington, N. E. Papua.

Leucophaeini

The Leucophaeini are considered to be the basic tribe of the
Epilamprinae and as such have a combination of features that are
modified in the more specialized groups. The tegmina, while moder-
ately broad, may be more acute apically than in most of the other
tribes. The subcosta has irregular and crowded anterior rami in the
apical portion. This limiting of rami is found also in the Panchlorini
and rarely in other groups. The radius has its anterior rami more
irregular than usual, but still less so than in the Oniscosomini. A
definite apical posterior branch is present and the cubital branches are
oblique as in most of the tribes. The anal field approximates in length
the subcosta.

The wings have a long subcosta as in most of the tribes, but Ri
has fewer anterior rami than in other members of the group except the
Calolamprini. The remainder of the radius is relatively simple, but
the apical posterior sector is much more distinct than in the other tribes
and is secondarily branched. The cubitus has almost one-half of its
branches going to the apex, much as in the Paratropini, and more than

JOHN W. H. REHN 69

in most of the other tribes. The first plical vein is straight apically,
not curved forward as in some Epilamprini, Thoracini and Paranau-
phoetini. The third plical vein is entirely free from the branched
axillary.

The tegmina have a large humeral area. The subcosta has a num-
ber of irregular anterior rami, except in the basal third, which show
considerable terminal twigging. It is slightly longer than the anal
area. The radius has a number of more or less irregular anterior
rami, some of which have secondary branches. There are a number of
apical rami and a well-differentiated, branched, posterior radial branch.
Basally, media is close to radius and apically it has three or more
branches, some of which are irregular and may become fused with
portions of the radius. The cubitus has a rather large number of
sectors which go obliquely toward the postero-apical margin. A
number of these show secondary branching. Basally, between this
system and the plical furrow, there is a veinless area. The plical fur-
row is arcuate. The anal field has a considerable number of veins, the
more median of which usually have branches.

The wings have the preaxillary area longer, more produced, than
the relatively small anal area which is not produced. The plical notch
is moderate. The subcosta is long and simple, usually extending al-
most to the apical third. Ri parallels the subcosta, beyond this it has
three short anterior rami. The remainder of radius is parallel to Ri,
below its rami branching in a regular manner. A distinct, branched,
apical posterior branch is present. The media is a simple, somewhat
arcuate vein. The cubitus is moderately well developed with oblique
sectors some of which show secondary branching. Usually more than
one-third, almost half, of these go to the apical margin. Basally, there
is some subcubital twigging. The first plical vein is free, distinct and
straight. The second plical vein varies from slightly more than one-
third to about two-thirds the wing length, and always ends in the first
plical. The third plical vein is free, not stalked on the branched axil-
lary, straight to the wing margin, posterior to the plical notch. The
interplical area is not expanded. The branched axillary is well de-
veloped, having four or five simple branches, and there are twelve or
more simple anal veins posterior to it. Intercalated veins are present
in the anal field and at times in the preaxillary area.

MEM. AMER. ENT. soc., 14.

70 CLASSIFICATION OF THE BLATTARIA

The following genera that have been examined may be placed in
this tribe: Leucophaea and Phortioeca.

Representatives figured: Leucophaea maderae (Fabricius) figs. 37,
102. [Det. J. A. G. Rehn, 1937.] This species is the genotype.
Male. Bitje, Ja River, Cameroons.

Phortioeca phoraspoides (Walker) figs. 36, 101. [Det. Hebard,
1920.] The genotype is Zetobora peruana Saussure [== Phortioeca
peruana (Saussure)]. Female. San Carlos, Costa Rica.

Litopeltini

This tribe is related to the Epilamprini and Panchlorini on the one
hand and to the Phoraspidini on the other. The tegmina have the
large humeral area lacking, in its basal portion, the venation that occurs
in several tribes. The subcostal rami are regular as inthe Perisphaerini
and Calolamprini, but posterior branches are present as in the Leuco-
phaeini and Epilamprini. The radius is developed much as in the
Epilamprini, but the apical posterior branch is not well differentiated,
as in the Panchlorini. The branches of media and cubitus attempt to
parallel the posterior margin as in the Panchlorini.

The wings have the anal area intermediate in its degree of pro-
duction. It is larger and longer than in the basic tribes, but not pro-
duced as far as the preaxillary lobe, as in the Panchlorini and Epil-
amprini. In this it is more nearly comparable to the condition found
in the Phoraspidini. Ri is long as in the Epilamprini, but it has only
a few incomplete, irregular anterior rami, a condition only approached
in some of the other tribes. The radial sector has limited apical rami,
and a definite apical posterior branch is not present, thus as in the
Panchlorini and Thoracini. The cubitus has approximately one-third
of its branches going to the apex, in this agreeing with the more de-
veloped tribes. The third plical vein is stalked on the branched axil-
lary as in the Epilamprini, but there are not as many axillary branches
as in that tribe.

The tegmina are relatively broad, with the anterior and posterior
margins weakly arcuate. The plical notch is indicated. The humeral
area is expanded. The subcosta is very heavy and has a short series
of regular anterior rami, apically. Branched posterior branches are
present. The radius has a relatively few, at times branched, anterior

JOHN W. H. REHN 71

rami. The apical rami are present and the apical posterior radial
branch is not well differentiated. Media rises close to the radius and
has three, long, simple branches. The cubitus divides into two main
stems, each of which subdivides. These branches and those of media
tend to parallel the posterior margin. The area near the plical notch
is filled with additional short branches of cubitus. The plical furrow
is strongly arcuate. The anal area, which is about equal in length to
the subcosta including its posterior branches, has numerous simple
arcuate veins.

The wings have a well-developed preaxillary lobe, and the anal lobe
is almost equally produced apically. A shallow concave area repre-
sents the plical notch. The subcosta is simple and long, reaching the
anterior margin beyond the middle. Ri reaches the margin at about
the apical fourth and has a few irregular anterior rami. The radial
sector parallels Ri and has branched, incomplete, apical rami. A defi-
nite apical posterior radial branch is not present. The media appar-
ently rises from the middle of the arculus and is a simple, sigmoid vein.
The cubitus has a series of oblique, arcuate branches, the more basal of
which are irregular. About one-third of these branches go to the
apical margin. The first plical vein is simple and straight. The sec-
ond plical vein is relatively long and its end is free. The third plical
vein is stalked basally on the branched axillary and is straight. A
small interplical area is present. The branched axillary has only one
definite branch and behind it there are at least ten simple anal veins.
Intercalated veins are found throughout the wing and some may be
present in the tegmina.

It would appear that the genus Derocalymma may also belong to
this tribe. The tegmina in this genus, for the most part, are very
dense and such venation as is evident can not be fully interpreted. The
wing, on the other hand, resembles that of Litopeltis in most respects,
differing only in the shorter subcosta, fewer branches of cubitus, and
short second plical vein. Not as many of the cubital branches propor-
tionately go to the apical margin and the second plical vein ends in
the first plical vein.

Representatives figured: Derocalymma versicolor (Burmeister)
figs. 35, 104. [Det. J. A. G. Rehn, 1932.] This species is the geno-
type. Male. Kuke Pan, South Africa.

MEM. AMER. ENT. soc., 14.

N
bo

CLASSIFICATION OF THE BLATTARIA

Litopeltis bispinosa (Saussure) figs. 34, 103. [Det. J. A. G.
Rehn, 1927.] This species is the genotype. Male, female. Panama
City, Panama.

Epilamprini

This distinctive tribe is one of the more highly developed members
of the subfamily. The tegmina are relatively broad and have an ex-
panded humeral area that is usually practically filled with the subcostal
rami. These are more regular and stronger than in the other tribes
except the Nauphoetini, which has a narrower humeral area. The
radius has long, anterior rami, more spaced than in the Leucophaeini
and less so than in the Nauphoetini and Panchlorini. The apical rami
are longer and more regular than in related tribes. The media and
cubitus have their branches tending to parallel the posterior margin
much as in the Panchlorini and some other tribes, but the branches are
more numerous. The anal area is more elongate than in most tribes,
resembling the Thoracini, and not as extreme as in the Paranauphoe-
tint.

The wings have the anal area produced, equal to the preaxillary
area as in the Nauphoetini and Panchlorini, but the plical notch is more
evident than in those groups. The subcosta is longer than in most
tribes except the Leucophaeini and Ri has only a few rami as in that
group. The radial sector has its apical rami more regular than in
most tribes, and a short apical posterior radial branch is differentiated
as in the Nauphoetini. The cubitus has less than a third of its
branches going to the apical margin as in the Nauphoetini and Pan-
chlorini. The first plical vein is curved forward apically as in the Nau-
phoetini, Paranauphoetini and Thoracini. The third plical vein is
stalked basally on the branched axillary, and after becoming free is
close to it. This plical vein curves down apically as in the Paranau-
phoetini and Phoraspidini and a small interplical field is present.
There are a larger number, twelve or more, of simple anal veins than in
the less-developed tribes.

The tegmina are rather broad and the humeral area is well de-
veloped. The subcosta is always heavy basally and has rather regular,
usually heavy, anterior rami, and a simple, thickened posterior branch
may be present. The radius has rather regularly spaced, long, anterior

JOHN W. H. REHN 73

rami which normally have a few secondary divisions. The apical rami
may have subdivisions, and the apical posterior radial branch, when
present, is simple, either short or long. Media basally is close to
radius, and has from one to four branches which may subdivide. In
Epilampra this vein is simple. The cubitus has rather regular
branches which usually are from the stem that parallels the plical fur-
row. These branches, which may subdivide, vary from oblique to at-
tempting to parallel the posterior margin. The plical furrow is
strongly and evenly arcuate. The anal field has a rather large number
of veins, the more median ones rarely branched.

The wings have the anal lobe produced, equal to the preaxillary
lobe. The subcosta is simple and long, reaching the margin beyond
the middle. Ri has three or four simple terminal branches. The
radial sector has rather regular, but as times secondarily branched
apical rami. The apical posterior radial branch is always simple and
may be very short. The media apparently rises from the top of the
arculus and is simple, usually arcuate. The cubitus has a considerable
number of oblique branches normally very regular and evenly spaced,
rarely somewhat irregular without basal twigging. All but three or
four of the main branches go to the plical fold. The first plical vein
apically curves forward. The second plical vein varies from one-third
to two-thirds the wing length and usually ends in the first plical vein.
The third plical vein is stalked basally on the branched axillary, then
parallels it. Apically it curves down and a small interplical field is
present. The branched axillary has three or four simple branches and
behind this there are at least a dozen simple anal veins. The anal
field always has intercalated veins and the preaxillary portion usually
does except in the genus Epilampra.

In addition to the forms studied, it is believed that a considerable
number of genera belong to this tribe.

Representatives figured: Epilampra abdomen-nigrum (De Geer)
figs. 39, 105. [Det. Hebard.] The genotype is Epilampra nebulosa
Burmeister. Male. Caparo, Trinidad.

Homalopteryx laminata Brunner figs. 38, 106. [Det. Hebard,
1920.] The genotype is Homalopteryx capucina Brunner. Male.
Caparo, Trinidad.

MEM. AMER. ENT. Soc., 14.

74 CLASSIFICATION OF THE BLATTARIA

Morphna plana (Burmeister) figs. 40, 109. [Det. Hebard, 1935.]
The genotype is Epilampra maculata Brunner [== Morphna maculata
(Brunner)]. Male. Mousakanda, Gammaduwa, Ceylon.

Nauphoetini

This tribe is most closely related to the Panchlorini, but the teg-
mina and wings are not as specialized as in that group. The tegmina
in shape are intermediate between those in the Panchlorini and the
Epilamprini. The humeral area is not expanded as in the Pan-
chlorini, but is filled with numerous subcostal rami. The radius is
much as in that tribe, but usually has the apical posterior branch well
defined. The branches of media and cubitus are more oblique, going
toward the postero-apical angle. The subcosta is shorter than the anal
area as in some of the less specialized tribes.

The wings have the anal lobe produced, equal to the preaxillary
one as in the Epilamprini. The subcosta is as in the Panchlorini and
Thoracini, while the radius is more as in the Epilamprini with a defi-
nite apical posterior radial branch. The cubitus has about one-third
of its branches going to the apex as in both the Epilamprini and Pan-
chlorini. The first plical vein may be straight or upcurved apically, the
former as in the Panchlorini and the latter as in the Epilamprini. The
third plical vein is free from the branched axillary and is straight as in
the Panchlorini. However, the branched axillary has at least two defi-
nite branches as in many of the more generalized tribes.

The tegmina have the anterior and posterior margins subparallel
and the apical margin is broadly rounded as in the Panchlorini. The
plical notch is not evident. The humeral area is not expanded and it
is filled with the irregular anterior branches of the subcosta. The
subcosta, which may have posterior branches, is shorter than the anal
area. The radius has a series of spaced anterior rami which may
branch secondarily. Branched apical rami are present and an apical
posterior radial branch is at least indicated. The media has either two
or three branches and may have terminal twigging. The cubitus has
a number of regular, oblique branches, which may subdivide, that go
toward the postero-apical angle. The plical furrow is evenly arcuate.
The elongate anal area has a number of veins, some of which may
branch.

JOHN W. H. REHN 75

The wings have the preaxillary area relatively large and the anal
lobe is as produced as that area. The plical notch is indicated by a
shallow concavity. The subcosta is simple and reaches the margin
near the middle. RR: is distinct and has regular spaced anterior rami.
The radial sector has a few apical rami, and a relatively short, simple
apical posterior radial branch is present. The media is either stalked
on the radial sector or apparently rises from the top of the arculus. It
is a sigmoid vein that may have terminal twigging. The cubitus has
at least twelve regular, simple branches, of which not more than one-
third go to the apical margin. The first plical vein is either straight or
curved forward apically. The second plical vein varies from about
one-third to two-thirds the wing length and ends in the first plical.
The third plical vein is free from the branched axillary and is straight
to the margin. A small interplical field is present, larger in those
forms that have the first plical curved forward. The branched axillary
has at least one definite branch, but it is not possible to tell if there are
more divisions. In all, at least thirteen veins reach the margin in the
anal area. Intercalated veins are present in the anal lobe and may be
found in the preaxillary one. At times, they are found in the tegmina.

It is believed that in addition to the forms studied, a considerable
number of genera may be referred to this tribe.

Representatives figured: Leurolestes pallidus (Burmeister) figs. 41,
111. [Det. Hebard, 1919.] This species is the genotype. Male.
Key West, Florida.

Nauphoeta cinerea (Olivier) figs. 47, 110. [Det. Hebard, 1928. ]
This species is the genotype. Female. Fort de Kock, Sumatra.

Panchlorini

This tribe is most closely related to the Nauphoetini. The teg-
mina are more slender than in most of the tribes. The spaced
branches of cubitus that attempt to parallel the posterior margin are
characteristic. The humeral area is not expanded and has only the
terminal subcostal rami, as in the Calolamprini and Litopeltini. The
anterior and apical rami are much as in the Epilamprini, but are more
spaced as in the Nauphoetini.

The wings have a full anal lobe, but it is not as produced as the pre-
axillary one, thus being intermediate between the condition found in

MEM. AMER. ENT. soc., 14.

76 CLASSIFICATION OF THE BLATTARIA

the Leucophaeini and that in the Epilamprini. The subcosta is short
as in the basic tribes, but Ri has a large series of regular rami, either
more or more regular rami than in any other tribe. An apical pos-
terior radial branch is absent. The cubitus does not have more than
one-third of its branches going to the apex, as in the Epilamprini and
Nauphoetini. The first plical vein is straight, and the third plical vein
is straight and free from the branched axillary as in the Nauphoetini.
It is not possible to determine the number of branches of the axillary,
a condition practically distinctive as it occurs elsewhere only in the
highly modified Thoracini and Paranauphoetini.

The tegmina are relatively slender with the anterior and posterior
margins subparallel. The apex is broadly rounded and the plical notch
is at least indicated. The humeral area is not expanded. The sub-
costa usually has some regular anterior rami apically and it may have
a series of posterior branches (Panchlora). The radius has spaced
anterior rami, some showing secondary branching. The apical rami
have regular subdivisions. The apical branch is simple or not differ-
entiated. The media has its base close to radius and has either two or
three simple branches. The cubitus which is very regular has branches
that rise from the stem that parallels the plical furrow, and these are
more or less parallel to the posterior margin. The plical furrow is
evenly arcuate. The anal area has numerous veins, some of which
may have branches.

The wings have a relatively large preaxillary area and a definite
plical notch. The subcosta is simple and reaches the anterior margin
near the middle. R: is well developed, with a regular series of an-
terior rami, connected apically in Panchlora. The radial sector has a
few apical rami, and a definite apical posterior branch is not present.
Media usually rises from the top of the arculus and is a simple, arcuate
_vein. The cubitus has a number, usually twelve, of regular oblique
branches of which about one-third go to the apical margin. The first
plical vein is simple and direct. The second plical vein is short, at
times ending in the first plical. The third plical vein is distinct from
the branched axillary and is straight to the margin behind the plical
notch. The axillary is strong, but it is not possible to determine if it
has branches. There are at least twelve simple veins that go to the
margin of the anal field. Intercalated veins are present in the anal
field and may be present in the preaxillary area.

JOHN W. H. REHN 77

It seems probable that many genera in addition to those studied be-
long to this tribe.

Representatives figured: Panchlora cubensis Saussure figs. 48, 108.
[Det. Hebard, 1919.] The genotype is Panchlora pulchella Bur-
meister —= Blatta quadripunctata Stoll [= Panchlora quadripunctata
(Stoll) ]. Male. Villavicencio, Colombia.

Pycnoscelus surinamensis (Linnaeus) figs. 42, 107. [Det.
Hebard, 1928.] The genotype is Pycnoscelus obscurus Scudder =
Blatta surinamensis Linnaeus [== Pycnoscelus surinamensis (Lin-
naeus)]. Female. Fort de Kock, Sumatra.

Thoracini

The members of this tribe are highly specialized Epilamprinae.
One feature that will separate this complex from all other Blattaria is
the large forward-curving anal field of the wing.

The tegmina resemble in some respects those of several of the other
tribes. Thorax has a produced humeral area, and the radius much as
in the Epilamprini, while Oryhaloa agrees in most respects with the
Paranauphoetini and in others with the Panchlorini. Features such as
the short subcosta and elongate, subangulate anal field are also shared
with some Pseudomopinae. There does not appear to be a combi-
nation of tegminal features that would allow recognition of this tribe.

The wings show a close relationship with the Paranauphoetini and
agree in many respects with those of other tribes. The apically nar-
rowed preaxillary lobe and the produced, forward-curving anal field
are distinctive. The long Ri with numerous short rami and the lack
of a definite apical posterior radial branch are features shared with the
Paranauphoetini and Panchlorini. The cubitus has at least some of its
basal branches fusing with the first plical vein as in the Paranauphoe-
tint. The curving forward of the first plical vein and its fusing
apically with portions of the cubital branches is distinctive. The third
plical vein does not become free from the branched axillary until after
its last branch, as in the Paranauphoetini. The forward curving of
this vein combination is distinctive. There is a small interplical field
as in the Paranauphoetini.

The tegmina are moderately elongate and vary from rather slender
to relatively broad. The anterior margin is weakly arcuate, the apical

MEM. AMER. ENT. soc., 14.

78 CLASSIFICATION OF THE BLATTARIA

margin roundly angulate and the posterior margin is arcuate to the
plical notch. This notch is not well developed. The humeral area
may or may not be enlarged. The subcosta is heavy and direct, either
simple or with apparently a few slender anterior rami. The radius has
a number of regular anterior rami and apical rami are present. An
apical posterior branch is always well developed and shows at least
some secondary divisions, these going to the extreme apex. The
media basally is close to the radius, then diverges and has at least two
well-developed branches. Basally, between it and cubitus is an area
which is free from venation. The cubitus rises near and for a distance
parallels the plical furrow. This vein has six or more major branches
that go obliquely toward the postero-apical margin. The plical fur-
row is strong, evenly arcuate in Thorax and subangulate apically in
Oxyhaloa. he anal area has a number of veins; those in the middle
of the field are oblique and show some branching. In general, those
in the anterior portion are arcuate, and more or less parallel the plical
furrow, while those in the posterior portion are arcuate and attempt to
parallel the posterior margin.

The wings have a small preaxillary area which is distinctly con-
stricted apically because of the anterior prolongation of the anal field.
The anal field is very large, prolonged anteriorly and produced with a
full fan. The anterior margin of the wing is weakly arcuate. The
very small apical margin is evenly rounded. A shallow but distinct
plical notch is present, followed by the evenly arcuate margin of the
anal fan. The subcosta is strong and parallels the anterior margin.
In Thorax, it is simple and ends slightly beyond the middle, while in
Oxyhaloa its tip is bifurcate and it ends at about the middle of the
wing. The radius has Ri distinct and more or less parallel to the sub-
costa. Past this vein it has numerous anterior rami, regular or irreg-
ular; and it ends at approximately the apical fifth. The radial sector is
strong with a few apical rami, and more or less parallels Ri. No defi-
nite apical posterior branch is preserved. The media apparently rises
either from the top or the middle of the arculus and is sigmoid. The
cubitus is well developed and has a number of oblique sectors which go
to the plical fold. Rarely, these show some secondary division. In
Oxyhaloa, all these sectors fuse terminally with the first plical vein,
from which they are indistinguishable, while in Thorax only the apical

JOHN W. H. REHN 79

ones are fused with the first plical. The first plical vein is prominent,
strongly curved forward apically. The second plical vein varies from
about one-fifth to one-half the wing length and always terminates in
the first plical. The third plical vein is fused for most of its length
with the branched axillary. Apically, the last anterior branch from
the axillary is considered to represent this vein. A small but distinct
interplical field is evident. There are apparently eight branches to the
well-developed branched axillary and these support approximately one-
half of the anal field. Behind this, there are at least eight simple anal
veins.

To this tribe belong the genera Thorax and Oxyhaloa. Also the
peculiar genus Phlebonotus would appear to be a member of this com-
plex. Notolampra is a member of this assemblage but may be dis-
tinguished by its much larger plical field. In this form, the third plical
vein curves posteriorly instead of paralleling the first.

Representatives figured: Oxyhaloa buprestoides (Saussure) figs.
43, 115. [Det. J. A. G. Rehn, 1922.] This species is the genotype.
Male. Baracoa, Cuba.

Thorax porcellana Saussure figs. 49, 114. [Det. Hebard, 1935.]
This species is the genotype. Male. Battaramulla, Wp., Ceylon.

Paranauphoetini

This distinctive tribe is most nearly related to the Thoracini, but
differs from it in many respects. The tegmina are proportionately
more elongate than in any of the other tribes and their sigmoid an-
terior margin is also distinctive. The plical furrow is arcuate and
marks off an elongate anal field, equal to about one-half the tegminal
length, longer than in any other tribe, as in the Panesthiidae. The
simple sigmoid subcosta is like that of Oryhaloa except it is heavier,
and the well-developed branched apical posterior radial branch is as in
that genus. The regular anterior and apical radial rami are as in the
Thoracini and some Epilamprini. The three branched media and
numerous oblique cubital branches are as in many of the Epilamprinae.

The wings have a relatively small preaxillary area and a full anal
area as produced as the preaxillary. A small interplical field is present
and the plical notch is absent. It agrees in the first feature with the

MEM. AMER. ENT. soc., 14.

80 CLASSIFICATION OF THE BLATTARIA

more highly developed tribes. The subcosta, radius and media are es-
sentially as in the Thoracini. The cubitus has its basal branches fus-
ing with the first plical vein as in that group, but its apical branches
fuse terminally and form an accessory vein parallel to the first plical.
Thus none of the branches go directly to the apex. This condition is
distinctive, but resembles that found in the Thoracini. The third
plical vein is stalked on the branched axillary as in that group, but does
not curve forward apically.

The tegmina are more dense basally than in the other tribes. The
anterior margin is weakly sigmoid, and the apical margin is strongly
rounded. The subcosta is simple, sinuate, very heavy and equal to
about one-half the length of the anal field. The radius which is very
heavy basally, has a series of spaced, regular anterior rami before the
origin of the radial sector. Beyond this Ri has additional spaced an-
terior and apical rami. The apical posterior radial branch, which
rises at approximately the apical third, has some secondary divisions.
The media is strong and before the mid-point divides into two stems,
one of these later subdividing just below the origin of the radial sector ;
no evidence of terminal twigging is present. The cubitus is strong and
has five or more branches, which go diagonally towards the postero-
apical angle. A few indirect basal branches are also present. The
plical furrow is arcuate, and reaches the posterior margin near the mid-
dle. The exceedingly long, lanceolate anal field has traces of a few
simple veins.

The wing is subtruncate apically and lacks a distinct plical notch.
The subcosta is simple and extends to about the middle of the wing.
Ri has a number of rather irregular anterior rami. The radial sector
and media are stalked basally; past this point the radial sector parallels
Ri, terminally it has two simple branches. The apical posterior branch
is not well differentiated. Media after becoming free is simple and
direct. The cubitus has its basal sectors passing to the plical furrow,
all but the apical three join the first plical vein. These three are joined
terminally to form an accessory vein which goes to the wing apex.
The first plical vein is strong and upcurved apically. The second plical
vein is relatively short, ending in the first plical. The third plical
vein is stalked on the branched axillary, rising after the last axillary
branch, and is down-curved apically. The curving first and third

JOHN W. H. REHN 81

plical veins bound a small interplical field, which is comparable to that
of other Epilamprinae and in a way to the intercalated triangle found
in certain of the Pseudomopinae. No trace of folding in this area is
noted. The anal area has thirteen simple anal veins counting the mem-
bers of the branched axillary; how many of these are branches of this
vein cannot be determined. The entire anal field has intercalated
veins, but these are absent in the preaxillary region.

Only the nominate genus, Paranauphoeta is placed in this tribe.

Representative figured: Paranauphoeta lyrata (Burmeister) figs.
45, 112. [Det. Hebard, 1922.] The genotype is Blatta circumdata
Haan [= Paranauphoeta circumdata (Haan)]. Male. Wuching, N.
W. Borneo.

Phoraspidini

This tribe, although very distinct, appears to be most closely re-
lated to the Epilamprini, Thoracini and Paranauphoetini. The almost
subtrigonal tegmina and the slender subcosta with simple, regular rami
apically are distinctive. Irregular radial rami and a simple apical pos-
terior branch show some resemblance to the condition found in the
Epilamprini. The oblique cubital branches represent the primitive
condition in the complex.

The wings have a short branched subcosta and lack a definite Ri,
both distinctive characteristics. The radial rami are more developed
than in other tribes. The cubitus, with less than a third of its branches
going to the apical margin, is a feature shared with the Epilamprini.
The third plical vein has its apex curving posteriorly as in some Epi-
lamprini, and Paranauphoetini and the Paratropini.

The tegmina are practically subtrigonal in outline, with a greatly
expanded humeral area and a narrowly rounded apex. ‘The subcosta
is slender and has four simple anterior rami in its apical half. The
radius has a series of irregular, and secondarily branched anterior and
apical rami. A simple, apical posterior branch is present. Media
divides into two long branches, one of which subdivides apically. The
cubitus has four simple, oblique branches which go to the posterior
margin. It is less developed than in any of the other tribes. The
plical furrow is subangulate apically, much as in some Pseudomopinae.
No plical notch is present. Traces of a few simple arcuate anal veins
are evident.

MEM. AMER. ENT. soc., 14.

82 CLASSIFICATION OF THE BLATTARIA

The wing has the anal field slightly produced, much as in the Calo-
lamprini, Epilamprini and Panchlorini. The subcosta is short as in
the Calolamprini, but it is forked terminally, a condition that is dis-
tinctive. Ri is not differentiated. The radius has spaced anterior
rami, which rarely subdivide. A few apical rami and a short simple
apical posterior branch are present. The media is simple, weakly sig-
moid. The cubitus has over a dozen oblique sectors of which about
one-third go to the apex. The first plical vein is simple, shghtly up-
curved apically. The second plical vein is long and free apically. The
third plical vein is stalked basally on the branched axillary, apically it
curves posteriorly. A small interplical field is present. There appear
to be three or four simple branches to the axillary. Thirteen veins
reach the margin in the anal area. Intercalated veins are found
throughout the wings.

Only the typical genus is included in this tribe.

Representative figured: Phoraspis picta (Drury) figs. 50, 113.
[Det. Hebard, 1922.] This species has been indicated as the genotype.
Male. Brazil.

Paratropini

This tribe does not appear to be closely related to any of the others,
but it may best be compared with the Leucophaeini. The tegminal
shape is as in that group, as is the development of the subcosta, which
is as long as the anal field. Also, definite posterior subcostal branches
are present as in that and some other tribes. The large number of ir-
regular, and secondarily branched anterior radial rami are distinctive,
although easily derived from the condition found in other tribes. It
is not possible to differentiate fully between radius, media and cubitus
as they appear to have a common origin. Whether an apical posterior
radial branch is present or not can not be determined. The media is
certainly branched, but the number of branches can not be ascertained.
This vein and cubitus both have their branches practically parallel to
the posterior margin as in the Panchlorini.

The wings are much like those of the Leucophaeini, but the pre-
axillary area has its apex more angulate. Rz is only forked terminally
and is shorter than in the other tribes. The branched axillary has its
branches more terminal than in other tribes.

JOHN W. H. REHN 83

The tegmina are broad with the apex rounded. A large humeral
area is present and it is filled with the anterior subcostal rami. This
vein is very heavy, with apical heavy rami, and at least two posterior
branches. There are numerous irregular anterior radial rami, many
of which branch secondarily. The branches of media and cubitus are
numerous and parallel the posterior margin, and some of them sub-
divide. The plical furrow is evenly arcuate and there is no plical notch.
Only a few irregular veins can be found in the anal field, less than in
other tribes.

The wings have a relatively large preaxillary area, and a smaller,
not produced anal field. The subcosta is simple and goes to the an-
terior margin at about the middle. Ri reaches the margin immedi-
ately after the subcosta and has only a terminal fork. The radius has
a number of secondarily branched anterior and apical rami. A very
short, simple apical posterior branch is present. The media is weakly
sigmoid and forks near the apex. The cubitus has more than a dozen
oblique branches of which one-half go to the apex. The first plical
vein is straight and simple. The second plical vein is long and its apex
is free. The third plical vein is remote from the branched axillary
and its apex curves posteriorly. The branched axillary has three
branches which originate near the apical third. Behind this are
thirteen simple anal veins.

Only the typical genus is placed in this tribe.

Representative figured: Paratropes phalerata (Serville) figs. 44,
120. [Det. Hebard, 1920.] The genotype is Paratropes lycoides
Serville. Male. Muzo, Boyaca, Colombia.

BRACHYCOLINAE

The unusual genera placed in this subfamily appear to be unique in
many respects. Their venation is for the most part more irregular
than is normal in the Blattaria. Moreover, the shape of the wing is
such, through reduction, that it appears these insects are not capable
of flight. The strikingly marked irregularly veined tegmina and the
wings with their proportionately small anal area, incapable of being
flattened, and amorphous venation will differentiate the Brachycolinae
from other groups.

MEM. AMER. ENT. soc., 14.

84 CLASSIFICATION OF THE BLATTARIA

In the tegmina, the subcosta may have some anterior rami as in
primitive Blattidae. The radius has a distinct Ri and a well-developed
apical posterior branch. In Hormetica, there are a large number of
branches. In these features, they resemble the Blattinae and to some
extent the Blaberinae, particularly in the tendency for anterior crowd-
ing. The media is well developed with a number of branches, thus re-
sembling the Nyctiborinae, except that they do not fill the apical mar-
gin. Cubitus has numerous oblique branches, resembling the more
primitive blattids. The strongly arcuate plical furrow shows a re-
semblance to the condition found in the Epilamprinae.

The wings resemble, at least in the radial area, the condition found
in the Blattinae. The retention of a branched media could show re-
lationship to this or other primitive groups. Cubitus with its freely
branching sectors mostly going to the apex, resembles the condition
found in some Epilamprinae and the Blaberinae. An archidictyon at
the base of cubitus is also found in the Blattinae and Blaberinae. The
first plical being involved apically with the terminal coalescing of the
cubital branches, as well as the third plical being stalked, in Brachycola,
for a long distance on the branched axillary, are conditions found also
in the Epilamprinae, particularly the Thoracini.

The tegmina are relatively short and broad, and have a network of
small irregular cells, so that the actual venation is partially obscured.
The humeral area is well developed and the anterior margin is weakly
sinuate or rounded. The apical margin is rounded and the posterior
margin is arcuate. A definite plical notch is evident. The subcosta
may have a number of irregular anterior rami. Basally this vein is
heavy and sinuate, while ventrally it is produced to form a tegminal
catch. The radial system presents either a relatively simple branched
design or a mass of amorphous branches. Rz is either simple or with
anterior rami. The sector has irregular anterior branches, many show-
ing secondary branching and with some indication of terminal twig-
ging. The media is strong, but hard to differentiate from the other
systems, with at least three major branches. Some secondary division
is present. The cubital area has a stem which divides into three major
branches, each of which has a number of secondary branches. These
are either straight, oblique or arcuate. The plical furrow is strong,
evenly arcuate, terminating at a well-developed notch. The anal area

on

JOHN W. H. REHN 8

has an indeterminate number of simple anal veins. In general, they
are more or less parallel to the posterior margin.

The wings of this subfamily show considerable diversity, and due
to their extreme individual irregularity and inconsistency only general
tendencies are noted. The relatively large preaxillary portion is in
contrast to the small anal fan. The presence of a definite plical notch
is in common with most of the primitive types. The anterior margin
is nearly straight, roundly passing into the apical margin. The mar-
gin of the anal field is broadly and evenly arcuate. The subcosta is
either simple or apparently fused with Ri, which has a variable num-
ber of rami. The radial system has an indeterminate number of ir-
regular branches, many of these secondarily dividing and all with some
evidence of terminal twigging. An apical posterior branch appears to
be present. The arculus is definite in Brachycola and probably is rep-
resented in Hormetica although it is not well differentiated. Media is
a strong, but irregularly branched vein. At times, some of its
branches become fused or at least connected with some of the cubital
sectors. Practically all its branches subdivide and some show apical
twigging. Cubitus has a variable number of sectors, usually somewhat
more than half of which go to the plical area and the remainder to the
wing apex, many having secondary branching. The first plical vein is
well developed although it does not always extend to the apex. At
times it apparently has accessory branches apically and becomes in-
volved with the fused terminal portion of the apical cubital sectors.
The second plical vein is either relatively short and ending free, or rep-
resented by a series of sections showing no definite connections. The
third plical vein is either stalked on the branched axillary or it is free
practically to its base. In all cases it reaches the wing margin behind
the plical notch. The branched axillary has three or four branches,
followed by a series of from eight to fourteen simple (or practically so)
anal veins. The branched axillary is not set off from the other veins.
The entire anal field usually has intercalated veins, and at times the
apical portion of the preaxillary area also has these veins.

The exact relationships of these forms will probably have to be as-
certained by features other than wing venation. In looking at the
tegmina and wing one is impressed by their irregularity and pseudo-
primitiveness, reminding the observer of the atavistic condition which
is sometimes noted in other Orthoptera.

MEM. AMER. ENT. soc., 14.

86 CLASSIFICATION OF THE BLATTARIA

Representatives figured: Brachycola tuberculata (Dalman) figs.
46, 1195 Det Hebard-19215|5 sithis) is) them eenotypesmHemale
Organ Mountains, Brazil.

Hormetica apolinari Hebard figs. 51, 116. [Det. Hebard, 1920. ]
The genotype is Hormetica laevigata Burmeister. Female. Villavi-
cencio, Colombia.

BLABERINAE

These large distinctive blattids seem to retain a number of primitive
characteristics. In general, their venation is simple although it has be-
come increased by the secondary division of many of the veins. The
large humeral area agrees with that found in some of the Epilamprinae
as does the simply and regularly branched media. The regular,
oblique branches of cubitus indicate an affinity to this same subfamily.
The tendency for the apparent anterior crowding of radius is char-
acteristic.

The anterior crowding of the radius in the wings is distinctive.
The single, branched media is a condition intermediate between the
primitive and specialized Blattidae. The regular, oblique, branches of
cubitus, about one-half of which go to the plical furrow, is a condition
also found in the Epilamprinae. The third plical vein, which is dis-
tinct from the branched axillary, is comparable to that found in the
Blattinae and pseudomopoids. The long, simple subcosta and the
branched axillary with only a few branches, and not well differentiated
from the remaining anal veins, again are features that agree with the
Epilamprinae. This subfamily appears to occupy a position between
the primitive Blattinae and the more highly specialized Epilamprinae.

The tegmina vary from relatively short and broad to rather long
and relatively slender. At least in the genus Archimandrita, the
former condition is characteristic of the female. The anterior margin
is evenly arcuate and the plical notch is scarcely indicated. The mar-
gin of the anal field is rounded angulate basally, the remainder being
practically straight. A well-developed humeral area is present. The
subcosta is strong and extends more or less obliquely to about the mid-
dle of the anterior margin. It may have regular anterior rami
throughout its length, or these may be crowded, irregular and only
present in the apical half. A forked or divided posterior branch is
present.. The radius is well developed, but gives the impression of

JOHN W. H. REHN 87

being crowded anteriorly. It has a number of rather irregular anterior
rami many of which show secondary branching. There are some
apical rami with numerous subdivisions. A distinct apical posterior
branch is present, and it usually has some secondary divisions, these
normally terminating at or near the extreme apex. The media is well
developed having four or five major branches which may have second-
ary divisions, all terminating at the apical margin. The cubitus nor-
mally divides into two major stems, each of which divides into at least
three, normally four or more branches which in turn may subdivide.
These branches in general go obliquely towards the posterior margin
with the exception of the basal one, which tends to parallel the plical
furrow and by subdivision completely fills the area between this furrow
and the more oblique sectors. The plical furrow is strong and usually
evenly arcuate. The anal field ranges from considerably shorter than,
to almost as long as, the subcosta, and it has a considerable number of
simple arcuate veins. These in general tend to parallel the plical fur-
row and a few of the most posterior veins show some terminal branch-
ing. The tegmina have at least indications of intercalated veins
throughout most of their area.

The wings have a relatively large preaxillary area and a moderate
to well-developed anal fan. The preaxillary area is considerably
broader in the genus Archimandrita than in Blaberus. The anterior
wing margin is nearly straight and the apical margin is either evenly
and broadly arcuate (Archimandrita), or rounded obtuse-angulate
(Blaberus). The plical notch is relatively shallow and the margin of
the anal field is evenly and broadly arcuate. The subcosta is simple
and parallels the anterior margin, passing at least the midpoint of the
wing, and in Archimandrita extending almost to the apical margin.
Ri usually becomes free from the remainder of radius near the base
and parallels the subcosta for a considerable distance, and may have
either a terminal fork or a few rami which pass anteriorly to the sub-
costa (Archimandrita). The radial sector basally more or less paral-
lels Ri and at about the midpoint some apical rami develop. These
often show secondary and further divisions. The apical posterior
branch of the radius is always distinct and has either terminal twigging
or secondary branching. The media apparently rises from the top or
the middle of the arculus and is arcuate, normally with a division into

MEM. AMER. ENT. Soc., 14.

88 CLASSIFICATION OF THE BLATTARIA

two branches which may have apical twigging. The arculus is promi-
nent and either transverse or somewhat oblique. The cubitus is well
developed and has numerous branches, these varying from fifteen to
approximately twenty with about one-half going to the apical margin
and the other half to the plical furrow. Some of them have secondary
division, but terminal twigging it not evident. The more apical ones
are inclined to be arcuate while the more basal ones are oblique. The
first plical vein is strong and direct. The second plical vein may be
simple and complete or simple and equal to about one-half the wing
length, with its apex free. The third plical vein is free basally, but
close to the branched axillary and goes directly to the apical margin,
rarely showing terminal twigging. A small interplical field may be
present. Basally the area on either side of the second plical shows
some irregular short accessory veins. It is believed that these have
been developed to strengthen the exceedingly large wing. The
branched axillary is prominent and has at least three major branches,
rarely showing some secondary division. Behind this there are at
least twelve simple anal veins, and some other short veins which do not
have connections with the basal bar, but may be stalked on an anal vein.
The entire anal field has strong, well-marked intercalated veins. In
the genus Archimandrita intercalated veins are present throughout the
wing except in the subcostal area, while in Blaberus these are not as
strongly indicated.

To this subfamily belong the genera Blaberus and Archimandnita.

Representatives figured: Archimandrita tessellata Rehn figs. 52,
117. [Det. Hebard, 1917.] The genotype is Blatta marmorata Stoll
[= Archimandrita marmorata (Stoll)]. Male. Colon, Panama.

Blaberus giganteus (Linnaeus) figs. 53, 118. [Det. Hebard,
LOW7/a his ispecies! ismithe cenoty peu: Malesm\vinzonmmboyacas
Colombia.

PSEUDOMOPOID COMPLEX

The pseudomopoid complex includes the four related subfamilies
(Pseudomopinae, Anaplectinae, Ectobiinae and Ceuthobiinae). All of
these have the tegmina with rather simple venation. The subcosta,
while primitively with a few rami, or forked, is usually simple, while
in other groups it often has numerous rami. Posterior branches of

JOHN W. H. REHN 89

the subcosta are usually not present, while in the other groups, with
the exception of the Blattinae, they are common. The radius has more
regular and more nearly vertical anterior rami than in the other groups.
The apical posterior branch is variable. Media, while primitively four
branched as in the Blattinae and epilamproids, may be fused with
cubitus, a condition only rarely occurring in other groups. The cubi-
tus when not fused with media has its branches oblique, or tending to
parallel the posterior margin as in the Nyctiborinae, except when it is
reduced to a simple stem. When fused with the media the sectors are
more oblique than in the other groups, passing toward the posterior
margin. The anal veins go diagonally across the field, not all uni-
formly curving as in the Blaberinae, or showing branching as in the
other epilamproids and Blattinae.

The wings have the subcosta shorter than in other groups and it
is simple. Usually Ri is distinct and it may be simple or with some
rami much as in the Blattinae, but it is not as long as in other groups,
or as branched as in the epilamproid complex. The anterior branches
of radius are more regular than in the other groups, but could easily
have been derived from the Blattinae type. Media is normally simple
as in most of the epilamproids. The cubitus usually has all its
branches going to the apex. There are rarely more than six branches,
often more reduced; this condition which is diagnostic of the group has
clearly evolved from that found in the Blattinae. An appendiculate
field, supported only by the third plical vein or an intercalated triangle,
is developed by some components and this is also distinctive. The
third plical has its base remote from the branched axillary, as in the
Blattinae and certain of the epilamproid complex. The axillary has
not more than five branches and does not support a large portion of the

anal field.

PSEUDOMOPINAE

The Pseudomopinae, while the dominant member of this complex,
are an assemblage of very diverse forms, but their tegmina have a dis-
tinctive combination of features. In most of the forms the subcosta is
simple as in the Ectobiinae, but a few of the primitive forms have some
rami as in the Anaplectinae and Ceuthobiinae. The radius has spaced,
regular rami and usually some branched apical rami, while in the three

MEM. AMER. ENT. soc., 14.

90 CLASSIFICATION OF THE BLATTARIA

related subfamilies the apical rami are usually not differentiated. In
some tribes a distinct posterior apical branch is present, as in the Ecto-
biinae, but it ends at the apex of the tegmina, while in the Ectobiinae it
goes posteriorly. Other tribes lack this branch and resemble the
Ceuthobiinae and Anaplectinae. The media may be branched or it
may be reduced to a single vein as in the Ceuthobiinae. At times it is
fused with cubitus as in the Anaplectinae and Ectobiinae. The cubi-
tus, when it is free from the media, has its branches oblique, to more or
less parallel to the posterior margin, as in the Ceuthobiinae. When it
is fused with media the branches go obliquely, or arcuately, towards the
posterior margin as in the Ectobiinae. The plical furrow may be
angulate apically, but in some forms it is evenly arcuate as in the other
subfamilies. The moderate number of anal veins are, more than in the
other subfamilies of the complex, in general arcuate.

The wings do not have an appendiculate field as in the Ectobiinae
and Anaplectinae, but, at times, have an intercalated triangle, supported
by the third plical vein. The small interplical field, which is some-
times present, is reminiscent of the condition found in certain of the
Epilamprinae. The margin of the branched axillary area is not greatly
produced as in the Ceuthobiinae. The radius often has a definite
apical posterior branch as in the Anaplectinae, but, at times, is un-
differentiated as in the Ectobiinae and Ceuthobiinae. The cubitus
usually has six or fewer branches which go to the apex. It is not re-
duced to a simple vein, except where the entire wing is reduced, as in
the other subfamilies. The main vein stems do not coalesce apically as
in the Ectobiinae. The first plical vein is either straight or upcurved,
apically, but does not fuse with the cubitus as in the Ectobiinae, but
may fuse with one of its branches, or is not lost as in the Anaplectinae.
The second plical vein is often relatively long, as in related subfamilies,
either free or fused apically with the first plical. The third plical vein
is either straight or down-curved apically, not supporting an appen-
diculate field as in the Anaplectinae and Ectobiinae. When an inter-
calated triangle is developed, the third plical is straight and serves as
its support. If an interplical field is present, this plical vein curves
downward and bounds the area posteriorly. The branched axillary is
remote from the plical veins and is distinct from the other anal veins,
as in the other members of the group. This vein usually has four or

JOHN W. H. REHN 91

five branches, not reduced to two or three branches as in most re-
lated subfamilies. Anal cross veins are usually numerous, not reduced
to one or two series as in the related subfamilies.

The tegmina are usually relatively long and slender, but are
broader and shorter in the Baltini, Euandroblattini and in those forms
showing reduction. The anterior margin which is usually weakly ar-
cuate may at times be almost straight or subsigmoid. Those forms
with long tegmina have the apex rounded, at times narrowly so, while
the forms with shorter tegmina may have the apex either broadly
rounded or subtruncate. The posterior margin is either almost
straight or weakly arcuate. The plical notch is always indicated, but
is never very strong. The margin of the anal field is either arcuate or
subangulate basally. The subcosta is heavy, usually weakly sinuate
and reaches the margin near the basal third, rarely near the middle
(Baltini). In the most primitive tribes a few simple anterior rami,
and rarely posterior rami (Parcoblatta), are present, but usually this
vein is simple. A distinct Ri is not developed, but there is a regular
series of spaced anterior rami, rarely thickened (Baltini). Simple,
branched apical rami are usually present and a distinct apical posterior
branch may or may not be developed. The media, when it is free from
the cubitus has usually three, but may have from two to five, major
branches ; terminal twigging is present only in the Parcoblattini. The
cubitus, when free from the media, often divides into two stems that
further subdivide. Primitively the branches go toward the postero-
apical angle, while in more specialized forms they attempt to parallel
the posterior margin. At times cubitus is apparently reduced to a
single oblique stem. When the media and cubitus are fused basally,
their branches are usually oblique. The plical furrow is definite, ar-
cuate, at times subangulate apically and reaches the margin at or be-
yond the end of the subcosta. The anal area has diagonal, simple
veins, more numerous in the primitive tribes. Intercalated veins may
or may not be present.

The wings have the preaxillary lobe moderately developed, while
the anal lobe shows considerable variation. In the primitive tribes it
is not produced, much as in the Blattinae; it enlarges progressively
until it is as produced apically as is the preaxillary lobe, comparable to

MEM. AMER. ENT. soc., 14.

92 CLASSIFICATION OF THE BLATTARIA

the condition found in some of the tribes of the Epilamprinae. In ad-
dition, it also increases in area as in that subfamily. The plical notch
is primitively present, but is lost when either an interplical field or an
intercalated triangle is developed. The subcosta is usually simple,
rarely with a terminal fork (Parcoblattini) or clubbed (Euphyllo-
dromia) ; at times it is stalked basally with Ri. It reaches the margin
between the basal third and the middle. Rx is distinct, varying from
simple to having a number of anterior rami. In the primitive tribes
it has the greater number of rami and is longer. The radial sector
has spaced, regular, simple anterior rami; at times these are clubbed.
Apical rami, either simple or branched, are always present. A dis-
tinct apical posterior branch which may be simple, branched or with
terminal twigging is usually present, but at times it is not differ-
entiated. The arculus is always strong, usually vertical and the media
usually apparently rises from its top. This vein is simple, straight to
subsigmoid. The cubitus primitively has a number of branches and
these are greatly reduced in the more specialized tribes. In some
tribes basal twigging, and some branches going to the first plical vein,
are present. In these cases about one-half or more of the branches go
to the apical margin. In the more advanced tribes there are from two
to five branches, all of which go to the apex. The first plical vein is
either straight or upcurved apically, at times fusing with one of the
cubital branches. The second plical vein varies from long to rather
short and its apex is usually free. The third plical vein is distinct
from the branched axillary and is either straight or downcurved api-
cally. The branched axillary is distinct from the other veins, and
usually has four and at times five branches. There are from seven to
more than twelve simple anal veins. At least two series of cross-
veins, usually more, are present in the anal area. Intercalated veins
may be present.

This subfamily may be divided into the following tribes: Parco-
blattini, Eustegastini, Euphyllodromim, Ischnopterini, Euandro-
blattini, Neoblattellini, Pseudomopini, Blattellini, Supellini, Symplocini
and Baltini.

JOHN W. H. REHN 93

BALTINI
BLATTELLINI
SYMPLOC INI
NEOBLATTELLINI
SUPELLINI
PSEUDOMOP INI
EUANDROBLATTINI

EUPHYLLODROMINI
EUSTEGASTINI

PARCOBLATTINI

ISCHNOPTERINI

ANCESTRAL STOCK

Diagram 5.—Probable relationship of the components of the Pseudomopinae.

Parcoblattini

This tribe appears to be the most primitive in the group, having a
number of features found in other subfamilies, but also having the
basic features which are further developed in the more modified tribes.
The tegmina retain the rami of the subcosta, a feature found elsewhere
only in the Ischnopterini, but common in the Blattinae and epilam-
proids. The radius has branched apical rami and a branched apical
posterior branch as in some Neoblattellini, Supellini and other tribes.
The distinct four-branched media is the basic type for the subfamily.
The cubitus is divided into two stems and has a number of oblique
sectors like those in the Ischnopterini. The plical furrow is angulate

MEM. AMER. ENT. soc., 14.

94 CLASSIFICATION OF THE BLATTARIA

apically or at least more strongly rounded apically, as in the Pseudo-
mopini, Symplocini and some other tribes.

The wing has the subcosta forked apically, a condition not normally
occurring in this subfamily. The radius is in general like that of the
Ischnopterini, except that there is a simple apical posterior branch.
The cubitus has six branches that go to the apex and some of these
branches secondarily divide, a unique condition in this subfamily.
There is some basal twigging as in the Eustegastini. The plical notch
is present and the plical veins are much as in the Euphyllodromini.

The tegmina vary from relatively long (male) to rather short
(female). The subcosta has spaced, regular rami in its apical half,
and a simple posterior branch may be present. The anterior radial
rami are rather regular, at times branched and the branched apical
rami may have terminal twigging. The branched apical posterior
branch shows terminal twigging. The distinct media has four long
branches which may show terminal twigging. The cubitus has five or
six simple branches that go to the postero-apical angle. The plical
furrow is strong, more angulate apically in the male. Some of the few
anal veins may rarely branch.

The wings have, for the subfamily, a relatively small anal lobe and
the plical notch is indicated. The subcosta reaches the margin at the
basal third and is forked terminally: Ri has a series of regular rami
reaching the margin in the middle third. The radial sector has
branched and simple anterior rami in addition to the branched apical
rami. A relatively short, simple apical posterior branch is present.
The media is stalked basally on the radius, and is sigmoid in its free
portion. The cubitus has a series of branches, almost all of which go
to the apical margin. Some secondary branching is present and in ad-
dition, there are a few short basal spurs. The first plical vein is not
complete and is slightly upcurved apically. The second plical vein is
straight and incomplete. The third plical vein is free and straight to
the margin behind the plical notch. The branched axillary is distinct
and has five branches, the more anterior of which may show terminal
twigging. In addition, there are about a dozen simple anal veins.

It is believed that a considerable number of genera may be placed
in this tribe.

on

JOHN W. H. REHN 9

Representative figured: Parcoblatta pensylvanica (De Geer) figs.
54, 55, 121. [Det. Hebard.] This species is the genotype. Male.
Havana, Illinois. Female. Ivy Hill, Mount Airy, Pennsylvania.

Eustegastini

The Eustegastini is a very distinctive group based on the typical
genus and is another tribe having strong Epilamprinae tendencies.
The tegmina have the radius much as in the Symplocini and Blattellini
except that it is sigmoid and the apical posterior branch is simple. The
media is free and three-branched. Anterior to this are two veins that
lack basal connections, but probably represent additional branches.
Thus, media may have more branches than usual for the subfamily and
these all go to the margin before the apex, a distinctive condition. The
cubitus has numerous branches rising at an angle, somewhat as in the
Pseudomopini, but the anterior stem has secondary branches instead
of being simple. The plical furrow is evenly arcuate as in some other
tribes, and the anal field has more veins than is usual in this subfamily.

The wings have a simple subcosta as in most of the complex. Ri
has the rami more numerous than in the other groups, but slightly
more so than in the Ischnopterini. The radial sector has only apical
rami, a distinctive condition. The cubitus has four branches going to
the apex and at least four short branches going to the plical fold, in
addition some basal stubs are present, somewhat as in Ischnopterini,
Parcoblattini and some Epilamprinae. The first plical vein is slightly
upcurved apically as in many tribes. A plical notch is present as in
the Parcoblattini and certain other groups, while the branched axillary
has only three branches. This vein is less distinct from the other anal
veins than is usual, which is much like the condition found in the
Ischnopterini.

The tegmina have a simple relatively heavy sigmoid subcosta. Ri
is not differentiated and there is a series of spaced, regular anterior
radial rami. The radius is definitely sigmoid and has a few apical
rami, more anterior than in other tribes. A relatively long simple
apical posterior branch is present. The media is strongly curved and
has three equal, simple branches. Two simple veins without basal con-
nections are anterior to it and probably represent additional branches.
All of these go to the margin before the extreme apex. The cubitus

MEM. AMER. ENT. Soc., 14.

96 CLASSIFICATION OF THE BLATTARIA

divides near the base and the anterior branch subdivides. The other
stem has a series of simple branches that rise at obtuse angles, and that
more or less parallel the posterior margin. The plical furrow is evenly
and strongly arcuate. A considerable number of simple anal veins are
present. Intercalated veins are present in the radial and cubital areas.

The wings have the anal lobe moderately developed, not produced
apically and the plical notch is evident. The subcosta is simple, sub-
parallel to the anterior margin which it reaches slightly before the
middle. Ri is long with ten simple, spaced rami and the radial sector
has a few simple apical rami and a relatively long, simple, apical pos-
terior branch. Media apparently rises from the top of the arculus and
is almost straight. The cubitus has four simple, relatively long
branches that go to the margin, then four shorter branches going
to, or almost to, the first plical and some short basal stubs. The
first plical vein is slightly upcurved apically. The second plical
vein is moderate in length and ends in the first plical. The third
plical vein is not far removed from the branched axillary, and is
slightly downcurved apically. There are apparently three simple
branches to the axillary, followed by nine simple veins. Intercalated
veins are present throughout the wing.

Only the typical genus is placed in this tribe.

Representative figured: Eustegasta poecila (Schaum) figs. 63, 127.
The genotype is Epilampra buprestoides Walker [= Eustegasta
buprestoides (Walker)]. Male. Amani, “German East Africa”
[= Tanganyika Territory].

Euphyllodromini

The Euphyllodromini are a small complex that resembles the Isch-
nopterini in many features. The tegmina have the radius and media
as in that group and the Euandroblattini, except that the media is re-
duced to two or three branches. The cubitus is from two to four
branched; these branches tend to parallel the posterior margin and are
somewhat angulate basally, reminiscent of the condition found in the
Euandroblattini.

In the wings, the subcosta is free or stalked on Rs, simple or
clubbed, much as in the Ischnopterini and Euandroblatta. Rx is simple
as in many of the specialized groups. The radial sector has regular

JOHN W. H. REHN 97

spaced rami which may be clubbed, as in the Supellini and some Neo-
blattellini. Apical rami and an undifferentiated apical posterior
branch are as in the latter tribe. The cubitus has two or three
branches as in the Supellini and Blattellini. The first plical vein is
straight or upcurved apically and the third is straight, agreeing on one
hand with the basic type and on the other with the Neoblattellini and
other forms with an incipient or actual intercalated triangle.

The tegmina have the subcosta heavy, straight or almost so, and
shorter than the anal field. Ri is not differentiated and there is a
series of regular, spaced, rami that continue to the apex. Rarely these
are branched. ‘The apical posterior branch is not differentiated. The
media has two or three long, simple branches that attempt to parallel
the posterior margin. The cubitus parallels the plical fold basally and
has either two or four simple branches that more or less parallel the
posterior margin. The plical furrow is weakly arcuate, at times sub-
angulate apically. A few simple anal veins pass diagonally across this
field.

The wings have moderate to well-developed anal lobes, in the latter
case produced as much as the preaxillary lobe. A small area, appar-
ently comparable to the intercalated triangle, is present in Chroma-
tonotus, but no trace of folding is noticeable. In Euphyllodromia this
area is not specialized. The subcosta may be free and clubbed, or, it
is stalked on Ri basally. R: is distinct and may be clubbed. The
radial sector has a series of spaced rami which may be clubbed; simple
apical rami are present and an apical posterior branch is not differ-
entiated. The media is stalked for a short distance on the radius and
its free portion is either straight or sigmoid. The cubitus has either
two or three simple branches that go to the apex. The first plical vein
is either incomplete or upcurved apically. The second plical vein is
long and free apically. The third plical vein is straight or nearly so.
The branched axillary is moderately close to the third plical vein and
has four simple branches. In the forms with an upcurved first plical
the terminal branch of the axillary is downcurved; these two veins
mark off an area equivalent to the intercalated triangle of some tribes.
At least ten simple anal veins are posterior to the branched axillary.

In addition to the genera studied, it seems probable that other
forms may be placed in this tribe.

MEM. AMER. ENT. soc., 14.

98 CLASSIFICATION OF THE BLATTARIA

Representatives figured: Chromatonotus lamprus Hebard figs. 56,
122. [Det. Hebard, Paratype.] This species is the genotype. Male.
Corozal, Canal Zone, Panama.

Euphyllodromia angustata (Burmeister) figs. 57, 123. [Det.
Hebard, 1919.] This species is the genotype. Female. Gatun,
Canal Zone, Panama.

Ischnopterini

The Ischnopterini are another primitive tribe that shows some re-
lationship to the Epilamprinae. The tegmina have the subcosta pro-
portionately shorter than in the other tribes and they have short an-
terior rami, which are also found in the Parcoblattini. The radial
rami are regular basally and irregular apically; no distinct apical rami
or apical posterior branch is present, resembling in those respects the
Euphyllodromini and to some extent the Pseudomopini. The media is
free and four branched as in the Parcoblattini, and the numerous
cubital sectors are arcuate as is that tribe and some Euphyllodromini
and Neoblattellini. The plical furrow is rounded as in the Neoblat-
tellini and some other tribes.

The wings somewhat resemble those of the Eustegastini and some
Epilamprinae. The subcosta is stalked on Ri as in some other tribes,
but Ri has more rami than any other tribe except the Eustegastini.
The radius is essentially as in the Euphyllodromini. The cubitus has
four branches which go to the apex and five which go to the plical fur-
row, much as in the Eustegastini, but lacking the basal stubs found in
that tribe. The first plical vein curves forward apically, and the third
plical curves posteriorly, marking off a small interplical field as in the
Supellini.

The tegmina are elongate and the plical notch is evident. The
subcosta is relatively short and has a few short, apical rami. Rx is
not differentiated and the radial sector has a series of regular, spaced,
usually simple rami that continue to the apex; no apical posterior
branch is present. The media has four long simple branches. The
cubitus divides regularly and has six branches that go diagonally to-
ward the postero-apical angle. The plical furrow is regularly arcuate.
The anal field has simple diagonal veins.

JOHN W. H. REHN 99

The wings have the anal lobe moderately produced, shorter than the
preaxillary lobe, and a small interplical field is present. The subcosta
is stalked on R:, its free portion is sinuate. Ri has a series of spaced
rami and it ends beyond the middle. The radial sector parallels Ri
and past it has anterior rami, the more apical ones branched, and a few
simple apical rami. An apical posterior branch is not differentiated.
The media is simple and sigmoid. The cubitus has four simple apical
branches that go to the margin; basally there are four branches that go
to the first plical area. The first plical vein is upcurved apically. The
second plical vein is relatively long, ending free before the curving of
the other plical veins. The third plical vein is but moderately removed
from the branched axillary and is downcurved apically. The branched
axillary has four long simple branches, and posterior to it are at least
a dozen simple anal veins.

Although only the typical genus is placed in this tribe at this time,
it is believed that a considerable number of genera are closely related
to it and may be placed in this group.

Representative figured: Ischnoptera rufa rufa (De Geer) figs. 62,
126. [Det. Hebard.] The genotype is J. morio Burmeister. Fe-
male. Barro Colorado, Canal Zone, Panama.

Euandroblattini

The Euandroblattini represents another distinctive, but primitive
type of Pseudomopinae. The tegmina are proportionately broader than
in any of the other tribes and their subtruncate apex is also distinctive.
The radius is much as in the Euphyllodromini and Ischnopterini, ex-
cept that the more apical rami are regularly divided. The media is
four branched as in the latter tribe and the Parcoblattini. The cubi-
tus has its few branches parallel to the posterior margin and angulate
basally, reminiscent of the condition found in the Symplocini and com-
parable to the cubital stem in Euphyllodroma. The plical furrow is
evenly arcuate as in most of the higher Pseudomopinae.

The wing has a much larger anal fan than in most of the other
tribes, but it is not produced apically, being shorter than the preaxillary
lobe. The radius is more like the primitive condition and that found
in the Epilamprinae, than like most Pseudomopinae, having fewer and
more irregular rami. The cubitus has five branches suggesting the

MEM. AMER. ENT. soc., 14.

100 CLASSIFICATION OF THE BLATTARIA

condition found in the Parcoblattini, but does not have the basal twig-
ging found in that tribe. A distinct plical notch is present and the
first and third plical veins are straight as in the Pseudomopini and
Parcoblattini, and the second plical vein is long and free.

The tegmina are proportionately short and relatively broad and
the apex is subtruncate. The plical notch is well developed. The
subcosta is shorter than the anal field, sinuate and heavy basad. Rx is
not differentiated, but the first radial ramus is close to the subcosta.
The radius has spaced, simple rami in its basal half, apically these and
the apical rami subdivide. A definite apical posterior branch is not
present. The media has four, long simple branches that practically
parallel the posterior margin. The cubitus basally parallels the plical
furrow, and then has four long branches that parallel the posterior mar-
gin. The plical furrow is strong, almost evenly arcuate. The anal
field has a number of simple diagonal veins.

The wings have a larger preaxillary and anal lobe than is usual in
the subfamily, but the latter is not produced apically. A shallow plical
notch is present. The subcosta is stalked on radius and Ri, but its
apex is free and simple. Ri after becoming free goes to the margin
beyond the middle; its apex is forked. The radial sector has a few
branched apical rami. A definite apical posterior branch is not present.
The media apparently rises from the top of the arculus and it 1s practi-
cally straight. The cubitus has five, long, simple, arcuate branches
that go to the apex. The first plical vein is almost complete, practically
straight. The second plical vein is relatively long and its apex is
free. The third plical vein is remote from the branched axillary and
is straight. The branched axillary has five, long, simple branches
and posterior to it are ten or more simple anal veins.

Only the typical genus is placed in this tribe.

Representative figured: Euandroblatta curta (Walker) figs. 61,
124. [Det. J. A. G. Rehn, 1937.] The genotype is Euandroblatta
propera Rehn. Male. Kisantu, Lower Congo District, Belgian
Congo.

JOHN W. H. REHN 101

Neoblattellini

The Neoblattellini show considerable diversity, but those forms
studied all seem to be representatives of a single stock. In the teg-
mina the subcosta is as long as the anal field, a condition found only in
the highly modified tribes. The radius is much like that of the Par-
coblattini and Euphyllodromini. This tribe resembles the Supellini
and Blattellini in having a distinct area without venation between the
radius, and the fused media and cubitus. The apical branches of this
latter combination continue to the apex, as in the Blattellini, and are
not oblique to the posterior margin as in the Supellini. The plical
furrow is somewhat angulate, but not as strongly so as in the primitive
Pseudomopinae.

The wings have Ri well differentiated, not branched as in the Balt-
ini and Euphyllodromini, but often clubbed. The radius has anterior
and apical rami and an apical posterior branch much as in the Parco-
Dlattini, but the anterior rami are often clubbed as in the Supellini and
some Euphyllodromini. The cubitus is four or five branched, thus
intermediate between the primitive Parcoblattini and Ischnopterini and
the advanced forms such as the Supellini, Symplocini and Pseudomo-
pini. The first plical vein curves forward apically as in some other
tribes, and the third plical is straight to the margin, often supporting an
intercalated triangle as in the Baltini. The base of the third plical
vein is rarely lost, a condition not noted elsewhere.

The tegmina are elongate and have the plical notch but moderately
indicated. The subcosta is almost straight, heavy and as long as the
anal field. Rx is not differentiated and the radial sector has spaced,
rarely branched anterior rami. A few branched apical rami are present
and the apical posterior branch is not well differentiated. The media
is fused with cubitus, so that no distinction can be made between their
four to eight major branches. The more terminal branches are
straight and go to the apex and the others may be arcuate and go to
this margin, or be oblique and go to the posterior one. Some second-
ary division and terminal twigging is noted. The plical furrow is
either arcuate or weakly angulate. A few simple, diagonal veins are
present in the anal field.

The wings have a moderately large anal lobe and the plical notch
may be indicated or not. An intercalated triangle is often present.

MEM. AMER. ENT. soc., 14.

102 CLASSIFICATION OF THE BLATTARIA

The subcosta is a simple vein that may be stalked on the radial system.
Ri is more or less parallel to the subcosta and may be clubbed. There
are a number of spaced, regular, often clubbed anterior radial rami.
Branched apical rami and a branched apical posterior radial branch are
present. The media is usually simple, rarely with a terminal fork, and
practically straight. The cubitus shows considerable variation, but has
four or five major branches that go to the apex. Rarely these
branches may subdivide or have terminal twigging and at times basal
stubs are present on the main stem. The first plical vein is upcurved
apically, at times fused with one of the cubital branches. The second
plical vein is relatively long and its apex is free. The third plical vein
is straight, rarely its base is subobsolete. This vein supports the inter-
calated triangle that is often present. The branched axillary is remote
from the third plical and has at least four branches. Fourteen or more
veins reach the margin of the full anal field.

In this tribe, all conditions from an expanded veinless area on
either side of the apex of the third plical vein to a definite intercalated
triangle are observed. The margin of the anal field bounding the
branched axillary may be either evenly arcuate or slightly produced.
This latter condition resembles, to some extent, that found in the Ceu-
thobiinae.

It is believed that in addition to the genera now placed in this
tribe, a considerable number of forms may be eventually included.

Representatives figured: Aglaopteryx gemma Hebard fig. 59a.
[Det. Hebard, Paratype.] This is the genotype. Female. Mobile,
Alabama.

Cariblatta delicatula (Guérin) figs. 58, 125. [Det. Hebard,
1924.| This is the genotype. Male. Montego Bay, Jamaica.

Latiblattella rehm Hebard figs. 60, 129. [Det. Hebard, Para-
type.]| This is the genotype. Male. Cocoanut Grove, Florida.

Neoblattella fratercula Hebard figs. 59, 128. [Det. Hebard.]
The genotype is Blatta adspersicollis Stal [== Neoblattella adspersi-
cols (Stal)]. Male. Lancetilla, Honduras.

Pseudomopini

The Pseudomopini, the typical tribe of the subfamily, are a distinct
entity showing both extreme specialization in some respects and, on the

JOHN W. H. REHN 103

other hand, a retention of primitive features. The tegmina have the
subcosta simple and the radius has only anterior rami as in the Euphyl-
lodromimi and Ischnopterini. The media is simple, arcuate and very
heavy based, a distinctive condition. Cubitus has a simple anterior
branch equivalent to one of the stems in other tribes, and the posterior
stem has five branches that are crowded basally and rise at an angle, a
condition not found in any other tribe. The Euandroblattini and
Eustegastini have a different type of an angulation in the cubital
branches. The plical furrow is angulate apically as in the Parcoblat-
tini, Blattellini and some other tribes. :

The wings have a moderately long, simple subcosta as in the Su-
pellini and Symplocini. R:z is distinct and with terminal forks as in
the Supellini and Blattellini. The radius is well developed, having
numerous regular anterior rami as in the Euphyllodromini and Blat-
tellini. A short apical posterior branch is present, but not well differ-
entiated. The cubitus has three branches as in the Supellini. The
third plical vein goes straight to the apex, and a definite plical notch is
present as in the Euandroblattini and Eustegastini.

The tegmina are elongate and slender, more so in the apical two-
thirds. The plical notch is evident. The subcosta is a little shorter
than the anal field, relatively heavy, simple and subangulate basally.
Ri is not differentiated. The radius is very heavy basally and is
strongly sigmoid, an apical posterior branch is not differentiated.
There is a complete series of regular, spaced, simple anterior rami, the
more proximal ones slightly thickened. The media, which is simple
and heavy basad, parallels the radius and is therefore sigmoid. The
cubitus which basally parallels the plical furrow has a remote, simple,
curved anterior branch that appears to represent the anterior stem.
There are five branches that rise at a strong angle from the posterior
stem. These branches become further apart apically and all go to the
tegminal apex. The plical furrow is strong, definitely angulate
apically. There are a small number of curved, simple anal veins.

The wings have a well-developed preaxillary lobe and a relatively
small anal lobe. The plical notch is very distinct. The subcosta is
relatively long, reaching the margin slightly before the middle, simple
and straight. Ra: parallels the subcosta and has a few simple terminal

MEM. AMER. ENT. soc., 14.

104 CLASSIFICATION OF THE BLATTARIA

rami. The radial sector has a series of regular, simple, spaced an-
terior and apical rami. An apical posterior branch is not differentiated.
The media rises from the apparent top of the arculus and is almost
straight. The cubitus has three long, simple, straight branches that
go to the apical margin. The first plical vein is straight and complete.
The second plical vein is relatively long and it ends in the first plical.
The third plical vein is straight to the plical notch. The branched
axillary is remote from the third plical and apparently has five simple
branches. More than ten veins reach the margin in the anal area.

Only the typical genus can be placed in this tribe at the present
time.

Representative figured: Pseudomops intercepta (Burmeister) figs.
64, 137. [Det. Hebard, 1926.] The genotype is Blatta oblongata
Linnaeus [== Pseudomops oblongata (Linnaeus)]|. Male. Cuerna-
vaca, Mexico.

Blattellini

The Blattellini is a distinct tribe that has an unusual combination
of features. The tegmina resemble those of the Neoblattellini by hav-
ing essentially the same type of radius, except that the apical posterior
branch is very long. The media and ctbitus are fused and possibly the
cubitus is reduced to the single posterior branch. This is much as in
the Baltini, except the branches go to the apex as in some Neoblattel-
lini, instead of to the posterior margin. The plical furrow is subangu-
late apically as in the Parcoblattini.

The wing has the subcosta and radius much as in the Symplocini
but the cubitus has only two branches, instead of three or more
branches as in other tribes, which is distinctive. The second plical
vein is shorter than in any of the other members of this subfamily, and
terminates in the first plical vein as in the Pseudomopini. The plical
notch is not definitely indicated and there is no trace of a special field
in this area, another distinctive combination of features.

The tegmina are relatively elongate and the subcosta, which is
shorter than the anal field, is heavy, simple and almost straight. Ri
is not differentiated and the radial sector has simple, spaced anterior
rami, the most apical of which may branch. A few apical rami, either
simple or branched, are present. There is a distinct apical posterior

JOHN W. H. REHN 105

branch that is equal to one-third of the tegminal length, and that has
terminal twigging. The media and cubitus are fused so that it is not
possible to separate their six long branches that go to the apex. The
most basal branch is remote, more or less paralleling the plical furrow
and may represent a single unbranched cubitus. The plical furrow is
distinct, subangulate apically. A very few simple veins, most of which
are oblique, are present in the anal area.

The wing has the preaxillary lobe relatively narrow and the anal
lobe is moderately produced, bounding the branched axillary. A
definite plical notch is not evident. The subcosta is simple and the dis-
tinct Ri has simple terminal rami. The radial sector has regular,
simple, spaced anterior and apical rami. The apical posterior radial
branch is equal to one-third the wing length and has terminal branches.
The media, which is fused for a short distance with the radius, is al-
most straight. The cubitus parallels the media and has two simple
apical branches that are but a little shorter than the apical posterior
radial branch. The first plical vein is weakly upcurved in its apical
portion. The second plical vein is shorter than the subcosta and ends
in the first plical. The third plical vein is straight. The branched
axillary is well removed from the third plical and has four, long, simple
branches. Posterior to those there are ten simple anal veins.

It would seem that a considerable number of forms may be referred
to this tribe when further studies are made.

Representative figured: Blattella germanica (Linnaeus) figs. 66,
136. [Det. Hebard.| This is the genotype. Male. Key West,
Florida.

Supellini

The Supellini are a distinct tribe of the Pseudomopinae, but show
some of the characteristics of the Ectobiinae. Forms varying from
fully alate to greatly reduced are members of this tribe. The discus-
sion is based fundamentally on the fully alate forms; those showing re-
duction have as a whole a simplification of the basic plan. The teg-
mina have regular anterior and variable apical radial rami, and the
apical posterior branch is much as in the Neoblattellini. The media
and cubitus are fused indistinguishably, with oblique branches all of
which go to the posterior margin, as in the Baltini, and differing from

MEM. AMER. ENT. Soc., 14.

106 CLASSIFICATION OF THE BLATTARIA

the Neoblattellini in which at least some of the branches go to the apex.
The very distinct space between the radius and the medio-cubital sys-
tem, with oblique sectors anterior and posterior to it, is a condition
only noted elsewhere in some of the Chorisoneurinae, but it is indicated
in the Baltini. The apically angulate plical furrow is much like that of
the primitive Pseudomopinae.

The wing has a relatively small anal lobe, and the plical notch is
not indicated, much as in Euphyllodromia and the Blattellini. There
is a long simple subcosta as in some Neoblattellini. Ri is more de-
veloped, as in the Blattellini, but has its rami clubbed as in some Neo-
blattellini. The radius has only a few anterior rami, less than in other
tribes, the apical rami and the apical posterior branch are much as in
the Parcoblattini and Pseudomopini. A three-branched cubitus, shows
a relationship with other forms, such as the Blattellini, Neoblattellini,
Euphyllodromini and Pseudomopini. The first plical curves forward
apically and the third curves posteriorly as in the Ischnopterini.

The tegmina are usually elongate, when reduced more or less
obovate. The plical notch is not noticeable. The subcosta is simple,
slightly curved and may be somewhat thickened. It is a little shorter
than the anal area. The radius does not have a distinct Ri, but has
regular, spaced, simple anterior rami. In those forms not showing re-
duction, simple or branched apical rami are present and there is a dis-
tinct, branched apical posterior branch. The media and cubitus are in-
distinguishably fused, leaving a narrow strip that goes from the base
to the apex without venation, between them and the radius. The fused
medio-cubitus has a number of regular, usually simple, oblique branches
that all go to the posterior margin. The most basal branch, in the re-
duced forms, is more or less parallel to the plical furrow. The plical
furrow is distinct and may be subangulate apically. There are a few
simple oblique veins in the anal field.

The wings have a moderately developed preaxillary lobe and a rela-
tively small anal lobe. The plical notch is not evident. The subcosta
is long and simple, reaching the margin near the middle. Ri more or
less parallels the subcosta and has clubbed apical rami. The radial
sector has a branched, clubbed anterior ramus, and then some simple
rami followed by a branched apical ramus. The apical posterior radial
branch is short and has a terminal fork. The media apparently rises

JOHN W. H. REHN 107

from the top of the arculus and is almost straight. The cubitus has
three simple almost straight branches that go to the apex. The first
plical vein curves forward apically. The second plical vein is of
moderate length and its end is free. The third plical vein is down-
curved apically. The apices of the first and third plical veins mark off
a small intercalated field, which is much like that found in some of the
Epilamprinae. The branched axillary is remote from the third plical
vein and usually has four simple branches. There are at least six
simple anal veins posterior to it.

It seems probable that a number of genera may be referred to this
tribe in the future.

Representatives figured: Ceratinoptera picta Brunner figs. 69, 140.
[Det. Hebard, 1920.] This is the genotype. Female. Gatun, Canal
Zone, Panama.

Supella supellectillum (Serville) figs. 67, 131. [Det. J. A. G.
Rehn, 1940.] This is the genotype. Male. Fort Leavenworth,
Kansas.

Symplocini

The Symplocini are another tribe that have a combination of dis-
tinctive features. The tegmina have the radius much as in the Blat-
tellini, but the apical posterior branch is more elongate, equal to more
than one-half the tegminal length, a distinctive feature. The anterior
radial rami are somewhat thickened as in the Baltini. The media is
free as in the primitive tribes and has three branches as in Chroma-
tonotus, while the cubitus divides into two stems with their branches
paralleling the posterior margin somewhat like the condition found in
Euphyllodroma and the Euandroblattini. The plical furrow is angu-
late-apical as in the Parcoblattini, Blattellini and Pseudomopini.

The wings have a large anal lobe, almost as full as that of the
Euandroblattini. The third plical vein is straight to the apex and
supports a small plical field, but it is not developed into an intercalated
triangle as in the Neoblattellini. The subcosta and radius are much as
in the Blattellini, but the apical posterior radial branch is longer. The
cubitus has four branches going to the apex and some basal twigging,
the latter resembling the condition found in the Parcoblattini while
the former is as in the Neoblattellini. The first plical vein curves for-
ward apically as in many tribes. There is an accessory vein between

MEM. AMER. ENT. soc., 14.

108 CLASSIFICATION OF THE BLATTARIA

the third plical vein and the posteriorly curved first branch of the axil-
lary, a condition not noted in any other members of the subfamily.

The tegmina are elongate and somewhat broader than is usual.
The plical notch is scarcely indicated. The subcosta is relatively heavy,
simple, straight and shorter than the anal field. Ri is not differ-
entiated and the radial sector has a series of simple spaced anterior
rami. A few mostly branched apical rami are present. “The apical
posterior radial branch, which has apical branches, is very distinct and
is equal to more than one-half the tegminal length. The media is
somewhat arcuate and has three simple branches. The cubitus divides
into two stems, the more anterior subdivides, and the posterior after
paralleling the plical furrow has three branches. The branches of
cubitus and media all parallel the posterior margin. The plical fur-
row is arcuate, subangulate apically. There are a few simple anal
veins.

The wings have the anal lobe large and as produced apically as the
preaxillary lobe. There is no plical notch. A small field is developed
around the apex of the third plical vein, but a definite intercalated tri-
angle is not present. The subcosta is simple, reaching the margin
slightly before the middle. Ri is very distinct; it parallels the sub-
costa and has a short series of simple spaced anterior rami. The radial
sector has a few simple anterior rami and branched apical rami. The
apical posterior radial branch is equal to one-half the wing length and
shows terminal twigging. The media rises from the apparent top of
the arculus and is weakly sigmoid. The cubitus more or less parallels
media and has four simple branches that go to the apex. The more
posterior of these branches are curved forward apically. Some short
stubs may be present on the cubital stem. The first plical vein is
strongly upcurved apically. The second plical vein is long, ending
just before the first plical curves, and its apex is free. The third plical
vein is straight. The branched axillary is remote from the third plical
and apparently has six simple branches, the most anterior of which
curves downward apically, while this branch and the first plical bound
a small field around the apex of the third plical vein. Apically be-
tween that vein and the axillary is an intercalated vein. Posterior to
the branched axillary there are at least a dozen simple veins.

It is not known how many other genera may be included in this
tribe.

JOHN W. H. REHN 109

Representative figured: Symploce capitata (Saussure) figs. 65,
130. [Det. Hebard.] This species is the genotype. Male. Havana,
Cuba.

Baltini

The Baltini are related to both the Neoblattellini and the Supellini,
all apparently being derived from a common ancestor, but are in some
ways more advanced than these other tribes. The tegmina have the
subcosta long as in these tribes, but the anterior rami of the radius are
weakly clubbed, which appears to be distinctive. The apical posterior
branch has secondary branches, stronger than in any of the other
tribes; but somewhat like those in Latzblattella and the Parcoblattini.
The media and cubitus are fused and their branches are oblique, an-
gling toward the postero-apical angle, somewhat as in the Supellini.
Intercalated veins are present throughout the tegmina, a condition not
common in other tribes.

The wings are much like those in the Neoblattellini except that the
radial rami are never clubbed, and the apical rami and apical posterior
branch are not as developed. The second plical vein goes at least to
the inner margin of the intercalated triangle, not shorter as in the Neo-
blattellini. The intercalated triangle is well developed and in folding
it crosses the first branch of the axillary, a distinctive feature. The
number of crossveins in the anal field is somewhat reduced, but not to
one or two series as in the forms with an appendiculate field.

The tegmina are shorter and broader than in the other tribes, and
gradually narrow apically. The plical notch is definite. The sub-
costa is simple, almost straight, and is as long as the anal field. Rx is
longer, more oblique and thickened than the anterior rami. The radial
sector has a series of spaced, usually simple, somewhat clubbed an-
terior rami. Branched apical rami, and a branched, long, apical pos-
terior branch are present. The media and cubitus are fused and have
six oblique branches that go to the posterior margin. The proximal
branch basally, parallels the plical furrow. A veinless area is between
the medio-cubitus and the radial system, but it is less distinct than in
the Supellini. The plical furrow is strong, definitely angulate apically.
The anal field has a few oblique, curved veins, the most posterior of
which is branched.

MEM. AMER. ENT. soc., 14.

110 CLASSIFICATION OF THE BLATTARIA

The wing has a large anal lobe that is as produced as the preaxil-
lary lobe. The plical notch is absent and there is a well-developed
intercalated triangle, which is bounded anteriorly by the first plical
vein and posteriorly goes to the area between the apex of the first and
second branches of the axillary. The apex of the third plical vein
forms most of its support, but the apex of the first branch of the axil-
lary, and at times the apex of the second plical vein, are in this field.
The subcosta is long and simple, reaching the margin slightly beyond
the middle. Ri is simple and closely parallels the subcosta. The
radial sector has a series of simple spaced anterior rami. Simple or
branched apical rami are present and the apical posterior branch is not
well differentiated. The media rises from the apparent top of the ar-
culus and is simple and straight. The cubitus has four long simple
branches that go to the apical margin. The apex of the most pos-
terior branch may fuse with the first plical vein which is upcurved api-
cally. The second plical vein is long, ending at or in the intercalated
triangle, its apex free. The third plical vein, which is remote from the
branched axillary, is straight and is the main support for the inter-
calated triangle. The branched axillary has four long simple branches.
Posterior to this are seven or more simple anal veins.

It is believed that additional genera may be placed in this distinc-
tive tribe.

Representative figured: Balta similis (Saussure) figs. 68, 138.
[Det. Hebard, 1925.] The genotype is Balta epilamproides Tepper.
Female. Kauai Island, Hawaii.

ECTOBIINAE

To this subfamily belong all those forms having an appendiculate
field supported only by the third plical vein, except the Anaplectinae.
In many respects, the group appears to be most closely related to the
Pseudomopinae. The tegmina have many features in common with
that group such as the fusion, at least at times, of media and cubitus so
that they cannot be differentiated. From the Anaplectinae, this sub-
family may be separated by having the subcosta simple and equal in
length to the anal field, and the less strongly arcuate plical furrow.
Also, the presence, except in the Plectopterini, of an apical posterior
radial branch, and the branches of media and cubitus going obliquely

JOHN W. H. REHN JUL

to the posterior margin, further differentiated this subfamily from the
other forms with an appendiculate field.

The wings while different from those of the Pseudomopinae show
certain basic similarities. All included genera have an appendiculate
field, usually set off by a transverse fold. The genus Ectobius has the
appendiculate field crumpled in repose and lacks the distinctive trans-
verse fold. This field is entirely supported by the third plical vein as
in the Anaplectinae. It seems that this might be a development from
some type such as Balta, which has an intercalated triangle supported
by the third plical vein. The wings lack an apical posterior radial
branch which is present in the Anaplectinae. The apical fusion of the
main veins to bound basally the appendiculate field, the more numerous
crossveins in the preaxillary portion of the wings, and the presence of a
complete first plical vein further differentiate this subfamily from the
Anaplectinae.

The tegmina are either elongate lanceolate or subtrigonal in shape.
The anterior margin is usually almost straight, at times somewhat
arcuate. The apical margin is evenly but narrowly rounded and the
posterior margin is practically straight. The plical notch is at most
subobsolete. The margin of the anal field is rounded angulate, more
so in Plectoptera than in other members of the group. The humeral
area is but moderately developed. The subcosta is a relatively heavy,
simple vein about equal in length to the anal field. It is usually almost
straight, rarely sigmoid. The radius is heavy basally and usually ex-
tends to the extreme apex. It has fairly regular, spaced, anterior rami,
some having secondary divisions. In all but the Plectopterini an apical
posterior branch or branches may be present. The development of this
feature varies considerably in individuals and in series of some species.
These branches may go to the tegminal apex, or to the posterior mar-
gin at about the apical third. The media and cubitus may be fused
basally so that it is not possible to separate them. When they can be
differentiated, the media is somewhat arcuate and has three or more
major branches. These curve toward the posterior margin and, at
times, have secondary branches. In these cases, the cubitus more or
less parallels media, and it may be simple or have up to four arcuate
branches that go to the posterior margin. Rarely a simple arcuate or
sinuate vein that goes to the posterior margin at about the middle of

MEM. AMER. ENT. Soc., 14.

112 CLASSIFICATION OF THE BLATTARIA

the tegmina appears to be the cubitus. In Plectoptera, media and cubi-
tus are a single stem which branches once near the middle of the teg-
mina. The more anterior branch goes to the apex and the other to the
posterior margin. The plical furrow usually is evenly arcuate but in
Plectoptera, it is almost straight. Usually, a trace of a plical notch is
present. The anal field has at most four arcuate veins, usually the
more anterior of these roughly parallel to the plical furrow, and the
posterior one is more or less parallel to the anal field margin.

The wing has the distinctive appendiculate field which, except in the
genus Ectobius, is separated by a transverse fold. In that genus, it
is represented by a distinctive area that does not materially alter the
wing outline. The anterior wing margin is usually sigmoid, concave
basally and convex apically. The apical margin is formed by the ap-
pendiculate field and may be rounded, subcircularly produced or
rounded subconical. In the latter case, the extreme apex is subobtuse
angulate, while in the former it is evenly arcuate. As this field is sup-
ported by the plical vein, no plical notch is present. The margin of
the anal field is regularly arcuate, occasionally somewhat angulate
basally. The subcosta is simple and equal to about one-third to one-
half the total wing length, reaching the anterior margin after the latter
has become convex. Apically, it may be forked. The radius is well
developed and extends to the edge of the appendiculate field. A num-
ber of anterior rami are present which are usually regular, rarely show-
ing some secondary branching. An apical posterior branch is not pres-
ent. The base of media is either not evident or this vein is stalked on
the radial system. Media is usually straight or gently arcuate, curving
forward apically toward the radial sector. The area between these two
veins is divided into regular or irregular cells by crossveins. The
arculus is strong, either inclined or vertical. Cubitus is unbranched
and somewhat sigmoid, and after passing the arculus is more or less
parallel to media. The cubitus is upcurved apically and joins the
media. The first plical vein rises near the base of cubitus, usually
curves posteriorly and then more or less parallels that vein. Apically
this vein curves forward to the cubitus. Thus, in all cases, the first
plical, cubitus, media and radius are joined or attempt to join apically
and form a portion of the inner boundary of the appendiculate field.
The second plical vein is strong, extending from slightly less than half

JOHN W. H. REHN 113

to almost two-thirds the wing length, and ends free. The third plical
vein is free basally, practically straight to the appendiculate field,
crosses the fold and continues to the apex of this field, except in
Ectobius. The entire appendiculate field is supported by this vein.
The branched axillary has its base distinct from the third plical and
normally has two or three branches. In all cases, these first two
branches join apically and delimit the inner margin of a portion of the
appendiculate field. In most cases, the third plical vein is joined to
the branched axillary by one or more crossveins. Behind this, there
are eight to eleven simple anal veins. Usually, there is a single series
of crossveins in the anal area. Rarely, these are absent, as in some
specimens of Chorisoneura, or they may be double in the anterior por-
tion of the field, a condition which has been noted in Ectobius.

In this subfamily after the anal fan has folded, fanwise, the wing
folds along a line anterior to or on part of, the third plical vein. Then
the appendiculate field folds over on the dorsal surface of the preaxil-
lary area. In Ectobius the appendiculate field is crumpled and lies in
the same relative position. In this subfamily, a definite transverse
fold, either vertical or oblique, usually separates the appendiculate field
from the remainder of the wing.

The three distinctive types have been examined in this subfamily
and they are believed to represent separate tribes. In much of the
older literature, these forms have been given subfamily or even higher
rank.

The following tribes are recognized: Ectobiini, Chorisoneurini and
Plectopterini.

Ectobiini

This tribe is erected for the typical genus and certain other genera
known to share its features. The tegmina are elongate, lanceolate as
in the Chorisoneurini, but have a sigmoid subcosta which is distinctive.
The radius normally has a number of posterior branches, some of
which pass to a considerable portion of the posterior margin, instead of
having these branches restricted and mostly going to the apical margin
(Chorisoneurini). The median and cubital systems are more oblique
and less sigmoid than in the other tribes, and the plical furrow is more
strongly arcuate. The anal field has only a few veins and these do not
closely parallel the margins. .

MEM. AMER. ENT. soc., 14.

114 CLASSIFICATION OF THE BLATTARIA

The wings may immediately be recognized by their lack of dis-
tinctive transverse fold. The appendiculate area is crumpled in re-
pose and forms a normally rounded wing apex, not produced as in the
other tribes. Moreover, the base of media is not preserved, while it is
present in the other tribes. The base of the third plical vein is not
preserved, and this vein after entering the appendiculate field distinctly
curves posteriorly ; these are again distinctive features. The branched
axillary has only two branches, instead of three, and these fuse apically
to limit the appendiculate field. Anteriorly in the anal field, behind
the branched axillary, a second series of crossveins may be evident, a
feature not found in the other tribes.

The tegmina are in general elongate oval with the humeral area
moderately developed. The subcosta is heavy, simple and sigmoid.
The radius basally is thickened and extends to the extreme apex; an-
teriorly throughout its course are a number of regular rami. In the
apical half, regularly spaced posterior branches are present, which pass
to the apical and posterior margins. Media is somewhat sigmoid and
has four simple branches which go to the posterior margin. The cubi-
tus appears as a single, simple sigmoid vein, it also going to the pos-
terior margin. The plical furrow is well indicated, but there is only
faint indication of the plical notch. Three simple anal veins are
present.

The appendiculate field of the wing is not produced, its apical mar-
gin broadly rounded. This field evenly joins the anal field; no plical
notch or angulation is evident. The subcosta is simple and slender,
curving forward to the anterior margin at or before the middle. The
radius is sigmoid and has regular anterior rami throughout its length,
no distinct Ri or apical posterior branch is present. The base of media
is lost and the remainder of the vein more or less parallels cubitus, it
is upturned apically, joins the radius, and is connected with the radial
sector by a number of irregular crossveins. The arculus is strong and
vertical. The cubitus basally is sigmoid, then parallels media and api-
cally curves forward to join that vein. Media and cubitus are joined
by a number of crossveins which divide this area into regular rectangu-
lar cells. The first plical vein rises near the cubitus and is sigmoid, it
curves forward apically to join the cubitus. These two veins are
joined by crossveins, which divide the area into subrectangulate cells.

JOHN W. H. REHN 115

Apically, the first plical, cubitus, media and radius are joined and de-
limit the inner margin of the anterior portion of the appendiculate field.
The second plical vein ends free and is equal to approximately one-half
the wing length. The third plical vein has its base lost. It is sigmoid
to the appendiculate field and in this field it curves posteriorly, going to
the margin behind the middle. The branched axillary goes almost to
the apical third, then branches once, terminally these branches fuse.
Several crossveins connect the third plical and the branched axillary.
These and the branched axillary bound the inner margin of the pos-
terior portion of the appendiculate field. Within the cell made by the
axillary branches are irregular crossveins. Eleven simple anal veins
are found posterior to the branched axillary. | Anteriorly, these are
connected by a double series of crossveins, then a single series and fi-
nally no crossveins.

Representative figured: Ectobius lapponicus (Linnaeus) figs. 70,
134. [Det. Saussure.] This species is the genotype. Male. Europe.

Chorisoneurini

To this tribe belong, in addition to the typical genus, a number of
related genera such as Mediastimia.

The tegmina have certain features such as the simple straight sub-
costa and the stem of radius not reaching the extreme tegminal apex, in
common with the Plectopterini, but the shape is like that of the Ecto-
bini. The anterior radial rami near the apex are secondarily divided,
which is distinctive of the group. The posterior branches which may
subdivide are usually but not always present, and go either to the apical
margin or the apical posterior angle. There is considerable variation
in the development of these branches. The media and cubitus may be
stalked basally as in the Plectopterini. There is considerable variation
in these two systems so that only certain generalizations can be made.
Whether they divide into two main stems or not, a number of branches,
more than in the other tribes, that curve towards the posterior margin
are present. When two stems are present, the media usually does not
have less than four branches, while the cubitus may have up to that
number. At times, all of these branches rise froma single stem. The
plical furrow is not strongly arcuate as in the Ectobiini. The plical
notch is more evident than in any of the other tribes. The anal area

MEM. AMER. ENT. soc., 14.

116 CLASSIFICATION OF THE BLATTARIA

has four simple veins, the more anterior paralleling the plical furrow,
while the most posterior parallels the posterior margin.

The anterior wing margin is gently arcuate, not sigmoid as in the
other tribes, and roundly passes into the margin of the appendiculate
field. This field is subcircular in outline, and not as produced as in the
Plectopterini. It meets the margin of the anal field at a distinct angle
as in that group. The subcosta is equal to less than one-half the wing
length and forks terminally, not simple as in the other tribes. The
radius more or less parallels the subcosta and goes to the basal margin
of appendiculate field. It has numerous anterior rami that are less
regular than in the other tribes. The media is stalked basally on
radius, resembling the condition found in the Plectopterini, and is al-
most straight to the apical third, then gently and arcuately curving an-
teriorly to join the radial system. The area between these two veins,
beyond the subcosta, is divided into subrectangulate cells by crossveins.
The arculus is strong and vertical. The cubitus is at first sigmoid,
then more or less parallels the media, at the apex it curves anteriorly
and joins the media. The first plical vein rises near the cubitus, curves
posteriorly and then is almost straight to the margin of the appendicu-
late field where it curves forward to join the cubitus. Apically, the
area between the cubitus and the first plical is divided into a few sub-
rectangulate cells by crossveins. The first plical, media and radius are
joined apically, and mark off the inner margin of the anterior portion
of the appendiculate field as in the Ectobiini. The second plical vein is
strong, reaching almost to the inner margin of the appendiculate field
and its end is free. The third plical vein is free basally as in the Plec-
topterini, practically straight to the appendiculate field, and then curves
forward slightly. Its apex is at the extreme apex of the appendiculate
field. The branched axillary is strong and has three branches as in the
Plectopterini. Apically, the first two join and form the inner bound-
ary of this portion of the appendiculate field. The most posterior
branch reaches to the margin of the anal field. Nine simple anal veins
are behind the branched axillary. In at least some instances, a cross-
vein connects the third plical with the branched axillary apically. At
times, a series of crossveins is found in the anal field.

Representative figured: Chorisoneura flavipennis (Saussure and
Zehntner) figs. 71, 135. [Det. Hebard, 1921.] The genotype is

JOHN W. H. REHN 130%

Blatta migrifrons Serville [= Chorisoneura nigrifrons (Serville) ].
Male, female. Venvidio, Sinaloa, Mexico.

Plectopterini

This is another tribe of the Ectobiinae, which in many respects is
very distinct.

The tegmina in general are elongate subtrigonal, not lanceolate as
in the other tribes. The subcosta is heavy, straight to the margin as in
the Chorisoneurini, but extends almost to the middle instead of being
shorter. The radius does not have any apical posterior branch as in
the Anaplectinae, and it seldom reaches the extreme apex. The media
and cubitus are fused basally, so that differentiation between them is
impossible as in some Chorisoneurini. The branches that are present
usually pass diagonally towards the apex, instead of to the posterior
margin. The plical furrow is almost straight, reaching the posterior
margin somewhat before the middle, instead of being arcuate.

The wings of these forms have the appendiculate field extremely
large, equal in length to that of the remainder of the wing. It is larger
than in the other tribes and is rounded conical in outline instead of
ovate. In general, the wing resembles that of the other tribes except
that crossveins are not present between media and cubitus, or between
cubitus and the first plical. The branches of the axillary do not fuse
apically to bound the appendiculate field as in the other tribes.

The tegmina are subtrigonal in outline. The anterior margin is
weakly arcuate, the apical margin is evenly rounded to obtuse angulate
and the posterior margin is straight. The plical notch is not evident.
The margin of the anal field is strongly arcuate. The humeral area is
relatively small. The subcosta is heavy, simple and goes to the an-
terior margin slightly before the middle. The radius is almost straight
and extends almost to the apex. It has a number of regular anterior
rami, but no apical posterior branches are present. The media and
cubitus are fused for a considerable portion of their length; usually a
single branch rises before the middle. These two sectors then pass
diagonally towards the apex, but actually go to the posterior margin.
There is a rather large area without venation between the posterior
branch and the plical furrow. The plical furrow is almost straight,

MEM. AMER. ENT. soc., 14.

118 CLASSIFICATION OF THE BLATTARIA

passing diagonally to the posterior margin which it reaches slightly be-
fore the middle. The anal area has traces of four simple veins. The
most anterior one parallels the plical fold and the most posterior one
more or less parallels the posterior margin. The other two which are
somewhat more basal in position are subparallel.

The wing has the tremendous appendiculate field distinctive of this
group. The anterior margin of the wing is sigmoid, and that of the
appendiculate field is strongly arcuate; the apex of this field is rounded
subobtuse angulate, and the posterior margin of the field is evenly ar-
cuate, joining the margin of the anal field at a distinct angle. In out-
line the appendiculate field is subconical. The margin of the anal field
is more or less evenly rounded. The subcosta is relatively long, nearly
straight, reaching the anterior margin at the base of the convexity. Ri
is distinct, parallels subcosta and is forked. The radius is strong,
paralleling subcosta, then almost straight to the transverse fold, termi-
nally upcurved. A number of regularly spaced anterior rami are
present, but there is no apical posterior branch. Media basally is
stalked on the radius, then almost straight to near the transverse fold,
where it curves forward to meet the radius. The area between the
radius and media is divided into a number of subrectangulate to sub-
quadrate cells, by crossveins. The arculus is strong and oblique. The
cubitus is practically straight to near the transverse fold, then curves
forward to meet the media. The first plical vein rises near the base of
cubitus, curves down, then parallels the second plical vein to near the
transverse fold, terminally curving forward. No crossveins are pres-
ent between media and cubitus or between cubitus and the first plical
vein. The first plical vein, cubitus, media and radius are joined api-
cally by their upturned tips, before the transverse fold, and form the
inner margin of this portion of the appendiculate field. The second
plical vein is strong, going almost to the transverse fold, its apex free.
The third plical vein is basally free, practically straight to the apex of
the appendiculate field. The branched axillary is arcuate and has three
branches, the anterior one curving down along the transverse fold.
The second branch meets the margin at the angulation formed by the
appendiculate field and the anal field margin. The third branch goes
directly to the margin of the anal field. Behind this are seven simple
anal veins. A crossvein connects the third plical and branched axillary

JOHN W. H. REHN 119

just before the transverse fold. A single series of crossveins is present
in the anal area.

Representative figured: Plectoptera porcellana (Saussure) figs. 72,
133. [Det. J. A. G. Rehn, 1922.] This species is the genotype.
Female. Cayamas, Cuba.

ANAPLECTINAE

The members of this subfamily, while superficially resembling the
Ectobiinae, differ from them in a number of important respects. The
subcosta is distinctive in having a few regular anterior rami, thus re-
sembling the Blattinae. The radius does not have any apical posterior
branch as in most of the Ectobiinae. Media is a three-branched vein,
with the branches attempting to parallel the posterior margin instead
of going to this margin, as in related forms. The first branch ends at
the extreme apex, and the other two at the apical portion of the pos-
terior margin. The cubitus appears as a simple, sigmoid vein ending
at the posterior margin. The notch at the end of the plical furrow is
more evident than in the Ectobiinae.

The wing, while having a large appendiculate field, is rather dis-
tinctive in that this is elongate oval, not like that of the Ectobiinae.
The subcosta is long as in related forms. The radius has a definite
apical posterior branch, a condition not found in the Ectobiinae, while
the base of media is lost as in some members of that subfamily. A
basal stub of the first plical is present, but the remainder of this vein is
completely lost, a condition not noticed elsewhere in the Blattaria.
Although the apex of all the major vein stems curve forward before the
transverse fold, they do not join apically as in the Ectobiinae. The
third plical vein supports the appendiculate field as in the Ectobiinae,
but reaches the margin posterior to the apex. The branches of the
axillary reach to the transverse fold, but do not fuse apically as in most
of the Ectobiinae.

In general, the tegmina are elongate oval. The anterior margin is
evenly arcuate, more so than in related forms. The apex is narrowly
rounded and the posterior margin is weakly arcuate. The plical notch
is evident. The margin of the anal field is evenly arcuate. The hu-
meral area is prominent. The subcosta is strong, direct, and reaches
the anterior margin a little before the middle. This vein has three regu-
larly spaced anterior rami. The radius is at first arcuate, then straight,

MEM. AMER. ENT. Soc., 14.

120 CLASSIFICATION OF THE BLATTARIA

reaching the apical margin slightly before the extreme apex. It has
approximately twelve regular, unbranched anterior rami. No apical
posterior branches are present. Media is apparently three-branched,
these more or less parallel to the posterior margin, the first branch go-
ing to the extreme apex, and the others to the posterior margin. The
cubitus is a simple sigmoid vein, and for over half its course parallels
the plical fold; its apex is at the posterior margin. The plical fold is
evenly and strongly arcuate, and a distinct plical notch is evident. The
four simple anal veins are more or less convergent towards the apical
angle of the field.

The wing has the appendiculate field equal to a little less than one-
third the total length. Folding is accomplished in the same manner as
in the Ectobiinae. The anterior wing margin is strongly sigmoid, and
the anterior margin of the appendiculate field is strongly arcuate.
Apically it is evenly arcuate and the posterior margin is arcuate.
The latter joins the margin of the anal field at an obtuse angle. The
margin of the anal field is evenly arcuate. The subcosta is long and
simple, reaching the margin at the start of the convexity. The radius
is sigmoid, with Ri differentiated as a more oblique, anterior vein con-
nected with the main stem by crossveins. There is a series of seven
anterior radial rami, and at about the middle of these anterior rami, a
posterior apical branch, which parallels the radius, originates. The
apical posterior branch of radius is connected to the remainder of the
radius by one crossvein. The base of media is not evident; this vein
is weakly sigmoid, more strongly so at the base and apex. It is con-
nected with the radius and the apical posterior branch by three cross-
veins, which divide the area into rectangulate cells. The arculus is
strong and oblique. The cubitus is sigmoid, roughly paralleling media,
and two crossveins connect these veins. Below the arculus is a stub
representing the first plical vein. No other trace of this vein can be
found. The second plical vein, which is approximate to the third pli-
cal, is sigmoid to the transverse fold. A single crossvein connects it
with the cubitus. The third plical vein is weakly sigmoid to the trans-
verse fold, then nearly straight to the margin of the appendiculate field.
The appendiculate field is supported only by the third plical vein. The
branched axillary is strong and has three major branches, one of which
subdivides. All of these branches except the most posterior end at the
transverse fold. The most posterior branch ends at the angle formed

JOHN W. H. REHN Ibe

by the margins of the appendiculate and anal fields. Behind this are
eight simple anal veins. No crossveins connect the plicals with one
another nor with the branched axillary, and there are no crossveins
between the members of the branched axillary. Behind this, there is
a single series of regular crossveins. The apices of all the systems an-
terior to the third plical vein curve forward, but they are not connected
apically. The branches of the axillary which reach the transverse fold
end blindly.

In this subfamily is placed Anaplecta and some related forms.

Although the tegmina and wings in these forms have achieved the
same general result as is found in Ectobiinae the details are sufficiently
different to warrant a separate subfamily.

Representative figured: Anaplecta lateralis Burmeister figs. 73,
132. [Det. Hebard, 1918.] This species is the genotype. Male.
Porto Bello, Panama.

CEUTHOBIINAE

This distinctive subfamily, with the tegmina superficially resem-
bling those of the Latindiinae, is clearly distinct from the Polyphagidae
and seems to be related on one hand to the Pseudomopinae and on the
other to the Ectobiinae and Anaplectinae. The tegmina have a forked
subcosta, a feature of primitive blattids; but they do not have any an-
terior rami, in this agreeing with some of the Pseudomopinae. The
radius has regular anterior rami basally as in that group, but the spaced
apical rami are distinctive. A simple media and a three-branched
cubitus, all paralleling the posterior margin, are again Pseudomopinae
features. However, the few spaced crossveins are distinctive. The
evenly arcuate plical furrow is reminiscent of the condition found in the
Ectobiinae.

The wing with the distinct lobe bounding the branched axillary
area, and as large as the preaxillary lobe immediately distinguishes
this group from all the other Blattaria. However, a slight tendency in
this direction is noted in some Epilamprinae. The simple subcosta —
and Ri with anterior rami are essentially Pseudomopinae character-
istics. The radius with a few anterior rami, a simple media and cubi-
tus all connected by spaced crossveins are characteristics shared with
the Ectobiinae and Anaplectinae. The lack of an apical posterior

MEM. AMER. ENT. soc., 14.

122 CLASSIFICATION OF THE BLATTARIA

branch of radius is a condition also found in the Ectobiinae. The
well-developed plical veins, and these being distinct from the branched
axillary, and the reduction in the number of crossveins in the anal field,
are shared with the forms that have an appendiculate field.

The tegmina are membranous and transparent, long and relatively
narrow. ‘The anterior margin is almost straight and passes into the
rounded apical margin. The posterior margin is practically straight.
There is no indication of a notch at the end of the plical furrow. The
humeral area is but moderately developed, and the subcosta is heavy
and short, being equal to about two-thirds the length of the anal area.
It has a strong terminal fork. The radial system has eight regularly
spaced anterior rami to slightly beyond the middle of the tegmina, be-
yond this there are a few rami dividing the marginal area into cells.
The media is a simple vein, close to the radius basally, then curving
away. It continues more or less straight to the extreme apex. Cubi-
tus rises close to the base of the media, curves down at first and divides
into two major branches, the more posterior of which again divides.
The three simple branches go to the apical margin. Both media and
cubitus parallel the posterior margin. The plical furrow is distinct and
evenly arcuate. Only traces of venation are evident in the anal field.
Irregular, spaced crossveins divide the area between the various vein
branches into a series of subrectangulate cells.

The wing has the anterior margin straight and the apical margin
rounded. A distinct plical notch is present. The margin bounding
the branched axillary is arcuately produced equal to the preaxillary
lobe; posterior to this is the evenly rounded margin of the remainder of
the anal field. The subcosta is simple, reaching the margin well before
the middle. Ri extends to about the middle and has a few (five) an-
terior rami. The radial sector extends almost to the apex of the wing
and has a number of irregular rami, some of which secondarily branch.
No apical posterior branch is present. The arculus is distinct and
oblique. Media, which is unbranched, is stalked basally for a short
distance on radius, then becomes free and goes to the wing apex. The
cubitus is another simple vein going sigmoidly to the apical margin.
The first plical vein is strong, simple, and slightly upcurved apically.
The second plical vein is about one-half the wing length, simple and
with its apex free. The third plical vein is free basally and extends to

JOHN W. H. REHN 123

the wing margin at the plical notch. The branched axillary has four
branches which go to the expanded margin. Posterior to it are six,
simple anal veins. Intercalated veins are present in this latter area.
The preaxillary portion of the wing, as well as that of the branched
axillary, is divided into a number of more or less rectangulate cells by
the irregular crossveins. Posterior to the branched axillary there is
one or more series of anal crossveins.

This distinctive subfamily contains only the genus Ceuthobia.

Representative figured: Ceuthobia fulvella (Rehn) figs. 74, 139.
[Det. Hebard, 1921.] The genotype is Ceuthobia lepta Hebard.
Male. Carcarana, Argentina.

OULOPTERYGIDAE

This distinctive family does not have any close relatives. The teg-
mina, with the dense punctation in the basal portion, the peculiar
venation and a faint plical furrow, do not closely resemble those of any
other members of the Blattaria. The subdiamond-shaped area, at
about the apical third, between the radius and media, which lacks
venation, is also distinctive.

The wing is very unusual with its appendiculate field lacking any
venation, and in being curled spirally in repose. Moreover, the large
area between radius and media which is divided into a number of ir-
regular cells, and the abrupt termination of all the plical veins at the
inner margin of the appendiculate field, will further differentiate this
family from any other known Blattaria.

The tegmina are very dense and have numerous punctures in
slightly more than the basal half; apically they become more membra-
nous and the venation is more readily visible. The anterior margin
curves forward basally making a large humeral area; beyond this it is
nearly straight to near the apex. The apical margin is broadly
rounded and the posterior margin is straight, the plical notch absent.
The subcosta is simple, very heavy basally and sinuate, reaching the an-
terior margin, beyond the basal third somewhat resembling the con-
dition found in some of the Pseudomopinae. The radius is strong and
straight in the basal half, having irregular, simple anterior rami similar
to some Pseudomopinae, beyond this it is curved forward to approxi-
mately the apical third, then recurved with its actual apex almost on a

MEM. AMER. ENT. soc., 14.

124 CLASSIFICATION OF THE BLATTARIA

line with its origin. Basally this portion has rather regular anterior
rami; after the anterior curve, rami are more irregular, often having
secondary branches, and going to the apico-anterior area. The media
is simple, parallel to the radius until latter curves anteriorly; at this
point, the media curves posteriorly and extends almost to the extreme
apex. A subdiamond-shaped area without venation is present between
these veins. Apically, they are connected by a series of crossveins.
The cubitus is at first close to media and radius, but becomes free be-
fore the basal third. It has four or more arcuate branches, some of
which divide, mostly ending at the apical margin, as in the Blattinae
and other primitive groups. The plical furrow is indicated by a weak
arcuate line going to the posterior margin slightly before the middle.
Apparently at least six simple unbranched anal veins are present; their
bases are not evident. Most of the tegmina are divided into a series of
irregular cells by the numerous crossveins. This is more pronounced
apically, where the tegmina are less dense.

The wing is divided into a relatively small preaxillary area, a mod-
erate sized appendiculate field, and a large, full anal area. The an-
terior margin is basally straight, between the middle and apical third
strongly arcuate, this arcuation roundly passing into the margin of the
appendiculate field. This field has its entire margin strongly arcuate.
The margin of the anal field is strongly and evenly arcuate. The sub-
costa is strong, simple and straight, reaching the costal margin at about
the midpoint, as in some Epilamprinae. Rz is parallel to its sector in
the basal half, then curves forward and ends in two simple branches.
The radial sector, after paralleling Ri, has a number of irregular an-
terior rami, some secondarily branched. A number of irregular cells
are formed in this area by crossveins. The media, which is simple,
apparently rises from the top of a small but distinct arculus, and is con-
cave throughout, apically ending at the radial sector much as in the
Ectobiinae. The area between it and radius is divided into a number
of oblique, transverse, irregular cells by crossveins, a condition dis-
tinctive of this family. The cubitus is strong and simple and parallels
media to the transverse fold, as in the Anaplectinae and Ectobiinae.
The simple media and cubitus represent a condition found in the other
groups with an appendiculate field. The first plical vein is strong and
distinct, slightly upcurved basally, then straight to the transverse fold.

JOHN W. H. REHN 12

ur

The second and third plical veins are well developed and end at the
transverse fold. The third is distinct from the branched axillary as in
the Blattinaeand Pseudomopinae. These veins are connected near their
apices and with adjacent veins by a series of apical crossveins. In the
Ectobiinae the plical veins are only connected with the adjacent veins.
The entire appendiculate field is without any trace of venation. The
branched axillary has an indefinite number of branches, as in some
Epilamprinae. Counting the branches of the axillary, there are
seventeen spaced, simple anal veins in the well-developed anal area.

In addition to Oulopteryx, it seems probable that Melyroidea Shel-
ford is a member of this family.

Representative figured: Oulopteryx meliponarum Hebard figs. 75,
141. [Det. Hebard, Paratype.] This species is the genotype. Fe-
male. Fazendo do Sobrado, near Passo Quatro, Minas Geraes, Brazil.

MEM. AMER. ENT. Soc., 14.

126 CLASSIFICATION OF THE BLATTARIA

OND

14.

15)

18.

19.

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de Zool., 2nd series, XIV, pp. 153-227.

1864. Mém. Hist. Nat. Mexique, 3 mémoire, pp. 16-28, 154-

169, text figs. 1-11, pl. I, figs. 1-4, 17, 18, 21, pl. Il, fig. 28.

. BRUNNER VON WaTTENWYL, K. 1865. Nouveau systéme des Blattaires,

xi + 426 pp., 13 pl.

. Waker, F. 1868. Catalogue of the specimens of Blattariae, in the col-

lection of the British Museum, London, 233 pp.

———. 1869. Supplement to the catalogue of Blattariae, catalogue of
the specimens of Dermaptera, Saltatoria and supplement to the Blat-
taria in the collection of the British Museum, London, pp. 119-156.

Saussure, H. pe. 1863. Mélanges Orthoptérologiques. Premiere Fas-
cicule, I, Blattides. Mém. Soc. Phys. Hist. nat. Genéve, XVII, pp.
17-171, 1 pl.

————. 1869. Mélanges Orthoptérologiques, Deuxieme Fascicule, II,
Blattides. Mém. Soc. Phys. Hist. nat. Genéve, XX, pp. 228-289, pl. 3.

20.

ai.

22.

28.

Zs).

30.

JOHN W. H. REHN WEE

1872. Mélanges Orthoptérologiques, Quatrieme Fascicule, WI
Blattides. Mém. Soc. Phys. Hist. nat. Genéve, XXIII. pp. 91-159,
pl. 10.

1893. Révision de la Tribu des Héterogamiens. Rev. Suisse
Zool., Genéve, I, pp. 289-318.

1895. Reévision de la tribu des Panesthiens et de celle des Epil-
ampriens (Orthoptéres de la Famille des Blattides). Rev. Suisse
Zool., Genéve, III, pp. 299-364, pl. 9.
and Zehntner, L. 1895. Révision de la tribu des Périsphaeriens

(Insecta Orthoptéres de la Famille des Blattides). Rev. Suisse Zool.,
Geneve, III, pp. 1-59, pl. 1.

BRUNNER VON WATTENWYL, K. 1882. Prodromus der Europaischen
Orthopteren, xxx + 266 pp., pls.

1893. Révision du systeéme des Orthoptéres et description des
especés rapportées par M. Leonardo Fea de Birmanie. Ann. Mus.
Stor. Nat. Genova, XXIII, pp. 5-230, pl. I-VI.

Kirpy, W. F. 1904. A synonymic catalogue of Orthoptera, vol. I,
Orthoptera Euplexoptera, Cursoria et Gressoria (Forficulidae, Hemi-
meridae, Blattidae, Mantidae, Phasmidae), 501 pp.

VERHOEFF, K. W. 1903. Uber die Nerven des Metacephalsegmentes und
die Insectenordung Oothecaria. Zool. Anz., 26, pp. 20-31.

. Navas, L. 1905. Mem. R. Acad. Cienc. y Artes, Barcelona, 3a epoca, V,

num. 13, p. 21.

. Krause, H. A. 1906. Insecktenborse, 23, p. 116.
. HanpiircH, A. 1903. Zur Phylogenie der Hexapoden. Sitz. Nat.

Klasse Akad. Wiss. (Wien), 112, pp. 716-738.
1906-1908. Die Fossilen Insecten und Phylogenie der Rezenten
Formen. ix + vi-+ 1430 pp., pls.

. SHELFORD, R. 1907. Ectobiinae. Genera Insectorum, 55, Orthoptera, 13

pp., 1 pl.
1908. Phyllodromiinae. Genera Insectorum, 73, Orthoptera,
29 pp., 2 pl.
————., 1910. Epilamprinae. Genera Insectorum, 101, Orthoptera, 21
DD, A Dl.

————. 1910. Blattinae. Genera Insectorum, 109, Orthoptera, 27 pp.,
2 pl.
REHN, J. A. G. 1903. Studies in American Blattidae. Trans. Am. Ent.
Soc., XXIX, pp. 259-290.
1904. Studies in Old World Forficulids or Earwigs, and
Blattids or Cockroaches. Proc. U. S. Nat. Mus., 27, pp. 539-560.
Cuoparp, L. 1922 Orthoptéres et Dermapteres. Faune de France, 3,
vi+ 212 pp., fig. Hee
Hanitscu, R. 1915. Malayan Blattidae. J. Straits Br. R. Asiatic Soc.,
69, pp. 17-178, 7 pl.

MEM. AMER. ENT. Soc., 14.

128 CLASSIFICATION OF THE BLATTARIA

39.

40.

41.

46.

47.

48.

49.

50.
51.

. Heparp, M. 1916. A new genus, Cariblatta, of the group Blattellites

(Orthoptera, Blattidae). Trans. Am. Ent. Soc., 42, pp. 147-186, 2 pl.

. Karny, H. H. 1921. Zur Systematik der Orthopteroiden Insecten.

Treubia, I, pp. 163-269.

. Hanpiirscu, A. 1925. In Schroder’s Handb. Ent., I], pp. 481-493.

1929. In Kukenthal, Handb. Zool., IV, pp. 821-839.

. Bruges, C. T. and Metanper, A. L. 1932. Classification of Insects. Bull.

Mus. Comp. Zool., 73, 672 pp.

. Forpes, W. T. M. (See reference No. 5.) :
. Bey-Brenxko, G. 1941. Blattoidea from Los Banos, Luzon, Philippines.

Philipp. J. Sci., 75, pp. 323-337, fig.

. Princis, K. 1946. Colombianische Blattodeen, gesammelt von Herrn

G. Dahl und Frau M. Althén-Dahl in den Jahren 1936-1939. K.
Fysiogr. Sallsk. i Lund Forh., XV1(16), pp. 144-158.

Bruiyninc, C. F. A. 1948. Studies on Malayan Blattidae. Zool.
Mededeelingen, xxix, pp. 1-174, ill.

Cuoparp, L. 1932. Un cas de microphtalmie lieé a l’atropie des ailes
chez une blatte cavernicole (Orthoptera). Livr. Cent. Soc. Ent.
France, p. 495.

Heparp, M. 1943. Australian Blattidae of the subfamilies Choriso-
neurinae and Ectobiinae (Orthoptera). Monogr. Acad. Nat. Sci.
Phila., no. 4, pp. 3, 9.

. Cuoparp, L. 1925. Le distribution géographique des Blattinae apteres

au subapteres (Orth. Blattidae). Ass. fr. Ar. Sci., Congres Liege
(1924), pp. 975-977.

ReuNn, J. A. G. 1932. On apterism and subapterism in the Blattinae
(Orthoptera, Blattidae). Ent. News, XLIII, pp. 201-206.

. Bruiyninc, C. F. A. 1948. Studies on Malayan Blattidae. Zool.

Mededeelingen, X XIX, pp. 2-34, fig. 1-25.

. CHoparp, L. 1922. Orthoptéres et Dermaptéres. Faune de France, 3,

pp. 3-4, 17-18, fig. 12.

. Heparp, M. 1916. A new genus, Cariblatta, of the group Blattellites

(Orthoptera, Blattidae). Trans. Am. Ent. Soc., 42, pp. 185, 186,
pl. XIII.

Suetrorp, R. 1907. Ectobiinae. Genera Insectorum, 55, Orthoptera,
hy 2 o (UY). Sie

Saussure, H. pe and ZEHNTNER, L. 1895. Histoire naturelle des Orthop-
téres. Hist. phys. nat. Madagascar, 23, pp. 1-147, pls. 1-5.
Saussure, H. ve. 1864. Mém. Hist. Nat. Mexique, 3 mémoire, pp. 16-
28, 154-169, text fig. 1-11, pl. I, fig. 1-4, 17, 18, 21, pl. II, fig. 28.
BRUNNER VON WatTeNwyt, K. 1865. Nouveau systéme des Blattaires,
pp. 9-12.

Comstock, J. H. 1918. The wings of insects, p. 125.

Renn, J. W. H. 1948. Permanent mounts of dissections to be kept with
pinned specimens. Ent. News, LIX, pp. 242-243.

Fig.

Fig.

JOHN WwW. H. REHN 129

EXPLANATION OF FIGURES
Plate I

6.—dArenivaga bolliana (Saussure) Tegmen. Male, Brownville, Texas.

7.—Heterogamodes roseni (Brancsik) Tegmen. Male, Bairam-Ali,
Transcaspia, U.S.S.R.

8.—Corydia petiveriana (Linnaeus) Tegmen. Male, Trichinopoly
Presidency, Madras, India.

9.—Homoeogamia mexicana (Burmeister) Tegmen. Male, Jalapa, V.C.,
Mexico.

. 10.—Tivia termes (Karny) Tegmen. Male, Vryburg, British Bechuan-

aland, South A frica.

. 11.—Holocompsa nitidula (Fabricius) Tegmen. Male, Lares, Puerto Rico.
ig. 12. Euthyrrhapha pacifica (Burmeister) Tegmen. Male, Mafaos, Brazil.
ig. 13—Eupolyphaga sinensis (Walker) Tegmen. Male, Ching-yang-fu,

Kansu, China.

. 14.—Polyphaga aecgyptiaca (Linnaeus) Tegmen. Male, Zakaki, Cyprus.
. 15.—Dyscologamia pilosa (Walker) Tegmen. Male, Grot by Baso,

Sumatra.

. 16.—Dyscologanua pilosa (Walker) Tegmen. Female, Fort de Kock,

Sumatra.

g. 17.—Latindia dohrniana (Saussure and Zehntner) Tegmen. Female,

Trinidad River, Panama.

. 18.—Compsodes delicatulus (Saussure and Zehntner) Tegmen. Male,

Cacao Trece Aguas, Guatemala.

Plate II

. 19.—Salganea morio (Burmeister) Tegmen. Female, Darjiling District,

Himalayas.

. 20.—Diploptera dytiscoides (Serville) Tegmen. Male, Honolulu, Hawaii.
. 21.—Periplancta americana (Linnaeus) Tegmen. Female, Trichinopoly,

Madras, India.

. 22.—Nyctibora noctivaga Rehn Tegmen. Male, Changuinola District,

Panama.

ig. 23.—Megaloblatta blaberoides (Walker) Tegmen. Male, Muzo, Boyaca,

Colombia.

_ 24,—Blatta orientalis Linnaeus Tegmen. Female, Bloomington, Indiana.
. 25.—Panesthia angustipennis (Illiger) Tegmen. Male, Java.

. 26.—Deropeltis erythrocephala (Fabricius) Tegmen. Male, Transvaal.
. 27.—Pseudoderopeltis brevicollis (Serville) Tegmen. Male, Pretoria,

Transvaal.
28.—Eurycotis floridana (Walker) Tegmen. Male, Biscayne Bay,

Florida.
29.—Blatta orientalis Linnaeus Tegmen. Male, Bloomington, Indiana.

MEM. AMER. ENT. soc.. 14.

130

CLASSIFICATION OF THE BLATTARIA

Flaten lle

. 30.—Calolampra trrorata (Fabricius) Tegmen. Male, “ Nouveau

Holland” [= Australia].

. 31.—Oniscosoma granicollis (Saussure) Tegmen. Male, Dunkeld, Vic-

toria, Australia.

. 32.—Blaptica dubia (Serville) Tegmen. Male, Buenos Aires, Argentina.
. 33.—Perisphaerus contiguus (Saussure) Tegmen. Male, Mount Lam-

ington, N. E. Papua.

. 34.—Litopeltis bispinosa (Saussure) Tegmen. Male, Panama City,

Panama.

. 35.—Derocalymma versicolor (Burmeister) Tegmen. Male, Kuke Pan,

South Africa.

g. 30.—Phortioeca phoraspoides (Walker) Tegmen. Female, San Carlos,

Costa Rica.

. 37.—Leucophaea maderae (Fabricius) Tegmen. Male, Bitje, Ja River,

Cameroons.

. 38—Homalopteryx laminata Brunner Tegmen. Male, Caparo, Trinidad.
. 39.—Epilampra abdomen-nigrum (DeGeer) Tegmen. Male, Caparo,

Trinidad.

. 40.—Morphna plana (Burmeister) Tegmen. Male, Mousakande, Gam-

maduwa, Ceylon.

Plate IV

. 41.—Leurolestes pallidus (Burmeister) Tegmen. Male, Key West,

Florida.

. 42.—Pycnoscelus surinamensis (Linnaeus) Tegmen. Female, Fort de

Kock, Sumatra.

. 43.—O-xryhaloa buprestoides (Saussure) Tegmen. Male, Baracoa, Cuba.
ig. 44.—Paratropes phalerata (Serville) Tegmen. Male, Muzo, Boyaca,

Colombia.

. 45.—Paranauphoeta lyrata (Burmeister) Tegmen. Male, Kuching, N. W.,

Borneo.

. 46.—Brachycola tuberculata (Dalman) Tegmen. Female, Organ Moun-

tains, Brazil.

. 47.—Nauphoeta cinerea (Olivier) Tegmen. Female, Fort de Kock,

Sumatra.

. 48.—Panchlora cubensis Saussure Tegmen. Male, Villavicencio, Colombia.
ig. 49.—Thorax porcellana Saussure Tegmen. Male, Battaramulla, W.P.,

Ceylon.

. 50.—Phoraspis picta (Drury) Tegmen. Male, Brazil.
ge. 51—Hormetica apolinari WHebard Tegmen. Female, Villavicencio,

Colombia.

Fig.
Fig.

Fig.
Fig.

Fig.
Fig.
Fig.
Fig.

JOHN W. H. REHN Hot

Plate V

g. 52.—Archimandrita tesellata Rehn Tegmen. Male, Colon, Panama.
ig. 53.—Blaberus giganteus (Linnaeus) Tegmen. Male, Muzo, Boyaca,

Colombia.

. 54.—Parcoblatta pensylvanica (DeGeer) Tegmen. Male, Havana,

Tllinois.

. 35.—Parcoblatta pensylvanica (DeGeer) Tegmen. Female, Ivy Hill,

Mount Airy, Pennsylvania.

. 56.—Chromatonotus lamprus Hebard Tegmen. Male, Corozal, Canal

Zone, Panama.

. 57.—Euphyllodromia angustata (Latrielle) Tegmen. Female, Gatun,

Canal Zone, Panama.

. 58.—Cariblatta delicatula (Guerin) Tegmen. Male, Montego Bay,

Jamaica.

. 59.—Neoblattella fratercula Hebard Tegmen. Male, Lancetilla, Honduras.
ig. 5JA.—Aglaopteryx gemma Hebard Tegmen. Female, Mobile, Alabama.

ig. 60.—Latiblattella rehni Hebard Tegmen. Male, Cocoanut Grove, Florida.
. 61.—Euandroblatta curta (Walker) Tegmen. Male, Kisantu, Lower

Congo District, Belgian Congo.

. 62.—Ischnoptera rufa rufa (DeGeer) Tegmen. Female, Barro Colorado,

Canal Zone, Panama.

Plate VI

ig. 63.—Eustegasta poecila (Schaum) Tegmen. Male, Amani, “ German

East Africa” [= Tanganyika Territory].

. 64.—Pseudomops intercepta (Burmeister) Tegmen. Male, Cuernavaca,

Mexico.

ig. 65—Symploce capitata (Saussure) Tegmen. Male, Havana, Cuba.
. 66.—Blattella germanica (Linnaeus) Tegmen. Male, Key West, Florida.

Fig.

67.—Supella supellectilium Serville Tegmen. Male, Fort Leavenworth,
Kansas.

68.—Balta similis (Saussure) Tegmen. Female, Kauai Island, Hawaii.

69.—Ceratinoptera picta Brunner Tegmen. Female, Gatun, Canal Zone,
Panama.

70.—Ectobius lapponicus (Linnaeus) Tegmen. Male, Europe.

71.—Chorisoneura flavipennis Saussure and Zehntner Tegmen. Male,
Venvidio, Sinaloa, Mexico.

72.—Plectoptera porcellana (Saussure) Tegmen. Female, Cayamas, Cuba.

73.—Anaplecta lateralis Burmeister Tegmen. Male, Porto Bello, Panama.

74.—Ceuthobia fulvella (Rehn) Tegmen. Male, Carcarana, Argentina.

75.—Oulopteryx meliponarum Hebard Tegmen. Female, Fazendo do
Sobrado, near Passo Quatro, Minas Geraes, Brazil.

MEM. AMER. ENT. soc., 14.

CLASSIFICATION OF THE BLATTARIA

Plate VII

. 76.—Arenivaga bolliana (Saussure) Wing. Male, Brownsville, Texas.
. 77.—Eupolyphaga sinensis (Walker) Wing. Male, Ching-yang-fu,

Kansu, China.

. 78.—Homoeogamia mexicana (Burmeister) Wing. Male, Jalapa, V.C.,

Mexico.

. 79.—Holocompsa nitidula (Fabricius) Wing. Male, Lares, Puerto Rico.
. 80.—Latinidia dohrniana (Saussure and Zehntner) Wing. Female, Trini-

dad River, Panama.

. 81.—Polyphaga aegyptica (Linnaeus) Wing. Male, Zakaki, Cyprus.
ig. 82.—Heterogamodes roseni (Brancsik) Wing. Male, Bairam-Ali,

Transcaspia, U.S.S.R.

. 83.—Dyscologamia pilosa (Walker) Wing. Male, Grot by Baso, Sumatra.
. 84.—Compsodes delicatulus (Saussure and Zehntner) Wing. Male,

Cacao Trece Aguas, Guatemala.

. 85.—Tivia termes (Karny) Wing. Male, Vryburg, British Bechuana-

land, South Africa.

. 86.—Euthyrrhapha pacifica (Burmeister) Wing. Male, Mafiaos, Brazil.
g. 87.—Dyscologamia pilosa (Walker) Wing. Female, Fort de Kock,

Sumatra.

Plate VIII

. 88.—Panesthia angustipennis (Iliger) Wing. Male, Java.
ig. 89.—Salganea morio (Burmeister) Wing. Female, Darjiling District,

Himalayas.

. 90.—Nyctibora noctivaga Rehn Wing. Male, Changuinola District,

Panama.

ig. 91—WMegaloblatta blaberoides (Walker) Wing. Male, Muzo, Boyaca,

Colombia.

ig. 92.—Diploptera dytiscoides (Serville) Wing. Male, Honolulu, Hawaii.
. 93.—Blatta orientalis Linnaeus Wing. Male, Bloomington, Indiana.
ig. 94.—Periplaneta americana (Linnaeus) Wing. Female, Trichinopoly,

Madras, India.

ig. 95.—Pseudoderopeltis brevicollis (Serville) Wing. Male, Pretoria,

Transvaal.

. 96.—Deropeltis erythrocephala (Fabricius) Wing. Male, Transvaal.

Plate IX

. 97 —Calolampra irrorata (Fabricius) Wing. Male, “ Nouveau Holland”

[= Australia].

. 98.—Oniscosoma granicollis (Saussure) Wing. Male, Dunkeld, Victoria,

Australia.

JOHN W. H. REHN 133

g. 99.—Blaptica dubia (Serville) Wing. Male, Buenos Aires, Argentina.
g. 100.—Perisphaerus contiguus (Saussure) Wing. Male, Mount Laming-

ton, N. E. Papua.

. 101.—Phortioeca phoraspoides (Walker) Wing. Female, San Carlos,

Costa Rica.

- 102.—Leucophaea maderae (Fabricius) Wing. Male, Bitje, Ja River,

Cameroons.

. 103.—Litopeltis bispinosa (Saussure) Wing. Male, Panama City, Panama.
. 104.—Derocalymma versicolor (Burmeister) Wing. Male, Kuke Pan,

South Africa.

Plate

. 105.—Epilampra abdomen-nigrum (DeGeer) Wing. Male, Caparo,

Trinidad.

. 106.—Homalopteryx laminata Brunner Wing. Male, Caparo, Trinidad.
- 107.—Pycnoscelus surinamensis (Linnaeus) Wing. Female, Fort de

Kock, Sumatra.

. 108.—Panchlora cubensis Saussure Wing. Male, Villavicencio, Colombia.
g. 109.—Morphna plana (Burmeister) Wing. Male, Mousakande, Gam-

maduwa, Ceylon.

. 110.—Nauphoeta cinerea (Olivier) Wing. Female, Fort de Kock, Sumatra.

. 111.—Leurolestes pallidus (Burmeister) Wing. Male, Key West, Florida.

. 112.—Paranauphoeta lyrata (Burmeister) Wing. Male, Kuching, N. W.,
Borneo.

. 113.—Phoraspis picta (Drury) Wing. Male, Brazil.

Plate XI

. 114—Thorax porcellana Saussure Wing. Male, Battaramulla, W. P.,
Ceylon.

. 115—O-xryhaloa buprestoides (Saussure) Wing. Male, Baracoa, Cuba.

. 116.—Hormetica apolinari WHebard Wing. Female, Villavicencio,
Colombia.

. 117.—Archimandrita tesellata Rehn Wing. Male, Colon, Panama.
. 118—Blaberus giganteus (Linnaeus) Wing. Male, Muzo, Boyaca,

Colombia.

. 119.—Brachycola tuberculata (Dalman) Wing. Female, Organ Moun-

tains, Brazil.

. 120.—Paratropes phalerata (Serville) Wing. Male, Muzo, Boyaca,

Colombia.

MEM. AMER. ENT. soc., 14.

134

CLASSIFICATION OF THE BLATTARIA

Plate XII

. 121.—Parcoblatta pensylvanica (DeGeer) Wing. Male, Havana, Illinois.
ig. 122.—Chromatonotus lamprus Hebard Wing. Male, Corozal, Canal Zone,

Panama.

. 123.—Euphyllodromia angustata (Latrielle) Wing. Female, Gatun, Canal

Zone, Panama.

. 124.—Euandroblatta curta (Walker) Wing. Male, Kisantu, Lower

Congo District, Belgian Congo.

. 125.—Cariblatta delicatula (Guerin) Wing. Male, Montego Bay, Jamaica.
g. 126.—Ischnoptera rufa rufa (DeGeer) Wing. Female, Barro Colorado,

Canal Zone, Panama.

. 127.—Eustegasta poecila (Schaum) Wing. Male, Amani, “ German East

Africa” [= Tanganyika Territory].

. 128.—Neoblattella fratercula Hebard Wing. Male, Lancetilla, Honduras.
g. 129.—Latiblattella rehni Hebard Wing. Male, Cocoanut Grove, Florida.

Plate XIII

g. 130.—Symploce capitata (Saussure) Wing. Male, Havana, Cuba.
g. 131.—Supella supellectilium Serville Wing. Male, Fort Leavenworth,

Kansas.

. 132.—Anaplecta lateralis Burmeister Wing. Male, Porto Bello, Panama.
. 133.—Plectoptera porcellana (Saussure) Wing. Female, Cayamas, Cuba.
ig. 134.—Ectobius lapponicus (Linnaeus) Wing. Male, Europe.

. 135.—Chorisoneura flavipennis Saussure and Zehntner Wing. Female,

Venvidio, Sinaloa, Mexico.

. 136.—Blattella germanica (Linnaeus) Wing. Male, Key West, Florida.
ig. 137.—Pseudomops intercepta (Burmeister) Wing. Male, Cuernavaca,

Mexico.

ig. 138.—Balta similis (Saussure) Wing. Female, Kauai Island, Hawaii.
ig. 139.—Ceuthobia fulvella (Rehn) Wing. Male, Carcarana, Argentina.
ig. 140.—Ceratinoptera picta Brunner Wing. Female, Gatun, Canal Zone,

Panama.

. 141.—Oulopteryx meliponarum Hebard Wing. Female, Fazendo do

Sobrado, near Passo Quatro, Minas Geraes, Brazil.

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INDEX

To the subfamilies, tribes, genera and species

abdomen-nigrum, Epilampra, 73
aegyptiaca, Polyphaga, 33
Aglaopteryx, 102

americana, Periplaneta, 54
Anaplecta, 121

Anaplectinae, 119

angustata, Euphyllodromia, 98
angustipennis, Panesthia, 47
apolinari, Hormetica, 86
Archimandrita, 88

Arenivaga, 33

Balta, 110

Baltini, 109

bispinosa, Litopeltis, 72
Blaberinae, 86

blaberoides, Megaloblatta, 58
Blaberus, 88

Blaptica, 66

Blatta, 53

Blattella, 105

Blattellini, 104

Blattidae, 47

Blattinae, 50

bolliana, Arenivaga, 33
Brachycola. 86
Brachycolinae, 83
brevicollis, Pseudoderopeltis, 54
buprestoides, Oxyhaloa, 79

Calolampra, 64
Calolamprini, 62
capitata, Symploce, 109
Cariblatta, 102

MEM. AMER. ENT. Soc., 14.

Ceratinoptera, 107
Ceuthobia, 123
Ceuthobiinae, 121
Chorisoneura, 116
Chorisoneurini, 115
Chromatonotus, 98
cinerea, Nauphoeta, 75
Compsodes, 39
Compsodini, 39
contiguus, Perisphaerus, 68
Corydia, 34

Corydiini, 33

cubensis, Panchlora, 77
curta, Euandroblatta, 100

dehrniana, Latindia, 39
delicatula, Cariblatta, 102
delicatulus, Compsodes, 29
Derocalymma, 71
Deropeltis, 53

Diploptera, 44
Diplopteridae, 42

dubia, Blaptica, 66
Dyscologamia, 35
dytiscoides, Diploptera, 44

Ectobiinae, 110

Ectobiini, 113

Ectobius, 115

Epilampra, 73

Epilamprinae, 59
Epilamprini, 72
erythrocephala, Deropeltis, 53
Euandroblatta, 100

il

Euandroblattini, 99
Euphyllodromia, 98
Euphyllodromini, 96
Eupolyphaga, 33
Eurycotis, 54
Eustegasta, 96
Eustegastini, 95
Euthyrrhapha, 42
Euthyrrhaphinae, 40

flavipennis, Chorisoneura, 116
floridana, Eurycotis, 54
fratercula, Neoblattella, 102
fulvella, Ceuthobia, 123

gemma, Aglaopteryx, 102
germanica, Blattella, 105
giganteus, Blaberus, 88
granicollis, Oniscosoma, 66

Heterogamodes, 33
Holocompsa, 40
Holocompsinae, 39
Homalopteryx, 73
Homoeogamia, 35
Hormetica, 86

intercepta, Pseudomops, 104
irrorata, Calolampra, 64
Ischnoptera, 99
Ischnopterini, 98

laminata, Homalopteryx, 73
lamprus, Chromatonotus, 98
lapponicus, Ectobius, 115
lateralis, Anaplecta, 121
Latiblattella, 102

Latindia, 39

Latindiinae, 36

Latindiini, 38

Leucophaea, 70
Leucophaeini, 68
Leurolestes, 75

Litopeltini, 70

INDEX

Litopeltis, 72
lyrata, Paranauphoeta, 81

maderae, Leucophaea, 70
Megaloblatta, 58 -
Megaloblattini, 57
meliponarum, Oulopteryx, 125
mexicana, Homoeogamia, 35
morio, Salganea, 47

Morphna, 74

Nauphoeta, 75
Nauphoetini, 74
Neoblattella, 102
Neoblattellini, 101
nitidula, Holocompsa, 40
noctivaga, Nyctibora, 57
Nyctibora, 57
Nyctiborinae, 54
Nyctiborini, 56

Oniscosoma, 66
Oniscosomini, 64
orientalis, Blatta, 53
Oulopterygidae, 123
Oulopteryx. 125
Oxyhaloa, 79

pacifica, Euthyrrhapha, 42
pallidus, Leurolestes, 75
Panchlora, 77
Panchlorini, 75

Panesthia, 47
Panesthiidae, 44
Paranauphoeta, 81
Paranauphoetini, 79
Paratropes, 83
Paratropini, 82
Parcoblatta, 95
Parcoblattini, 93
pensylvanica, Parcoblatta, °5
Periplaneta, 54
Perisphaerini, 66
Perisphaerus, 68

petiveriana, Corydia, 34
phalerata, Paratropes, 83
Phoraspidini, 81
Phoraspis, 82
phoraspoides. Phortioeca, 70
Phortioeca, 70
picta, Ceratinoptera, 107
Phoraspis, 82
pilosa, Dyscologamia, 35
plana, Morphna, 74
Plectoptera, 119
Plectopterini, 117
poecila, Eustegasta, 96
Polyphaga, 33
Polyphagidae, 25
Polyphaginae, 30
Polyphagini, 31
porcellana, Plectoptera, 119
Thorax, 79
Pseudoderopeltis, 54
Pseudomopinae, 89
Pseudomopini, 102
Pseudomops, 104
Pycnoscelus, 77

MEM. AMER. ENT. soc., 14.

INDEX

rehni, Latiblattella, 102
roseni, Heterogamodes, 33
rufa, Ischnoptera, 99

Salganea, 47

similis, Balta, 110

sinensis, Eupolyphaga, 33
Supella, 107

supellectillum, Supella, 107
Supellini, 105

surinamensis, Pycnoscelus, 77
Symploce, 109

Symplocini, 107

termes, Tivia. 36

tessellata, Archimandrita, 88
Thoracini, 77

Thorax, 79

Tivia, 36

Tiviinae, 35

tuberculata, Brachyccla. 86

versicolor, Derocalymma, 71

111

—

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MeEMorrs OF THE AMERICAN ENTOMOLOGICAL SOCIETY

An irregular serial, containing monographic papers by students of
authority in their respective subjects.

No. 1. The Cresson Types of Hymenoptera. By Ezra T. Cresson.
1916. 141 pp. Price $3.00. No. 2. The Blattidae of North America,
North of the Mexican Boundary. By Morgan Hebard. 1917. 284
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vision of the Dipterous Family Rhagionidae (Leptidae) in the United
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Price $4.50. No. 8. The Eumastacinae of Southern Mexico and Cen-
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$5.00. No. 13. Elachistidae of North America (Microlepidoptera).
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Classification of the Blattaria as Indicated by their Wings (Orthop-

tera). 1951. 134 pp., 13 pls. Price $5.00.

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