A new species of Zorotypus from central Amazonia, Brazil (Zoraptera, Zorotypidae)

2006·0.66 MB·English

by  Rafael, José Albertino

A tamerican museum
Novitates
PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024
Number 3528, 11 pp., 14 figures, 3 tables September 08, 2006
A New Species of Zorotypus from Central Amazonia,
Brazil (Zoraptera: Zorotypidae)
JOSE ALBERTINO RAFAEL1 AND MICHAEL S. ENGEL2
ABSTRACT
A new species of zorapteran (Polyneoptera: Zorotypidae: Zorotypus) is described and figured
from alates and apterous individuals captured in Central Amazonia, Brazil. Zorotypus amazonensis
Rafael and Engel, new species, is distinguished from related neotropical species by the setation of
the male vertex, the peculiar development of the male cerci, female metafemoral ornamentation,
female sternites, and presence of a true Cu vein in the hind wing base (permitting for the first time
an accurate homologization of hind wing veins in Zoraptera). The Brazilian zorapteran fauna is
briefly summarized and the homology and phylogenetic implications of cepahlic and wing
venational traits are discussed. The Embiodea + Zoraptera clade is dubbed mirorder Mystroptera
Engel.
INTRODUCTION 2005). Zorapterans are enigmatic, largely
ignored insects that have mystified entomolo¬
With only 33 living and six fossil species, gists since their discovery nearly a century
the Zoraptera are the least diverse orders of ago. While presently considered an order
insects3 (Engel, 2003a; Grimaldi and Engel, of Polyneoptera (Anartioptera: Plecopterida)
1 Instituto Nacional de Pesquisas da Amazonia, INPA, Caixa Postal 478, 69011-970 Manaus, Amazonas, Brazil
([email protected]).
"Division of Invertebrate Zoology, American Museum of Natural History; Division of Entomology (Paleoentomology),
Natural History Museum, and Department of Ecology & Evolutionary Biology, Dyche Hall, 1345 Jayhawk Boulevard,
University of Kansas, Lawrence, KS 66045-7163 ([email protected]).
3The Notoptera (consisting of Grylloblattodea and Mantophasmatodea) are the second smallest order, presently with 41
living species (vide Grimaldi and Engel, 2005: treated as separate orders therein but actually representing a single lineage
[Engel and Grimaldi, 2004; Grimaldi and Engel, 2005; Arillo and Engel, in press]). The recognition of Mantophasmatodea
as a separate order renders Grylloblattodea paraphyletic, as the various Mesozoic grylloblattodean fossils are basal to
modern Grylloblattodea + Mantophasmatodea. It is therefore only valid to consider the Mantophasmatodea as a suborder
(Arillo and Engel, in press).
Copyright © American Museum of Natural History 2006 ISSN 0003-0082
2 AMERICAN MUSEUM NOVITATES NO. 3528
and most closely allied to the webspinners4 TABLE 1
(Embiodea) (Engel and Grimaldi, 2000; Hierarchical Classification of Order Zoraptera
Grimaldi and Engel, 2005), the order has been
diversely interpreted and at times considered Family Zorotypidae Silvestri
as related to various polyneopterous orders Genus fXenozorotypus Engel & Grimaldi
(e.g., Kukalova-Peck and Peck, 1993; f X. burmiticus Engel & Grimaldi
Genus Zorotypus Silvestri
Yoshizawa and Johnson, 2005), within or
Incertae sedis
sister to Paraneoptera (e.g., Hennig, 1969),
|Z. cretatus Engel & Grimaldi
sister to Eumetabola (e.g., Beutel and Gorb,
Subgenus | Octozoros Engel
2001) , or even as sister to Holometabola fZ. acanthothorax Engel & Grimaldi
(Rasnitsyn, 1998), thereby spanning all major tZ. nascimbenei Engel & Grimaldi
clades of neopteran insects. Diagnoses of Subgenus Zorotypus Silvestri s.str.
supraspecific taxa can be found in Engel Z. amazonensis Rafael & Engel, n.sp.
(2003b, 2004) and Engel and Grimaldi (2000, Z. barberi Gurney
2002) and a preliminary phylogenetic hypoth¬ Z. brasiliensis Silvestri
Z. buxtoni Karny
esis in Engel (2003b) (refer to table 1 for the
Z. caudelli Karny
current hierarchical classification).
Z. ceylonicus Silvestri
Adults are dimorphic, occurring as winged
Z. congensis Ryn-Tournel
and eyed individuals that are usually brown to Z. cramptoni Gurney
black and are the dispersive forms of each Z. delamarei Paulian
species, while the apterous and blind morphs tZ. goeleti Engel & Grimaldi
are slightly paler in color and are the pre¬ Z. guineensis Silvestri
dominant type encountered. Nymphs are Z. gurneyi Choe
similar to the adults but lack pigmentation Z. hamiltoni New
Z. hubbardi Caudell
and lack spiniform setae on the metafemora.
Z. huxleyi Bolivar y Pieltain & Coronado G.
The order is principally pantropical in distri¬
Z. javanicus Silvestri
bution, and geographic records are largely
Z. juninensis Engel
limited to the type localities, with few species Z. lawrencei New
being documented for more than one country Z. leleupi Weidner
(a result of poor sampling rather than actual Z. longicercatus Caudell
evidence of high endemism, vide infra). Z. manni Caudell
The order is relatively ancient, with rela¬ Z. medoensis Elwang
Z. mexicanus Bolivar y Pieltain
tively modern species reaching back into the
Z. neotropicus Silvestri
Early Cretaceous (Engel and Grimaldi, 2002),
Z. newi (Chao & Chen)
a situation identical for its sister, the
fZ. palaeus Poinar
Embiodea (Engel and Grimaldi, 2006). As
Z. philippinensis Gurney
such, it will likely be from early fossils that the Z. sechellensis Zompro
greatest insights into resolving the placement Z. shannoni Gurney
of the highly autapomorphic zorapterans will Z. silvestrii Karny
be found. Unfortunately, pre-Cretaceous fos¬ Z. sinensis Hwang
sils of these small, relatively soft-bodied Z. snyderi Caudell
Z. swezeyi Caudell
insects will be a challenge to locate given the
Z. vinsoni Paulian
delicate conditions necessary for their suitable
Z. weidneri New
preservation.
Z. zimmermani Gurney
While the fossil record of Zoraptera is
sparse, so too is our knowledge of the living
Gurney, and Z. gurneyi Choe. Indeed, the
species in which most of our information
order is so little understood that many
derives from three North American species:
seasoned entomologists have never seen zor¬
Zorotypus hubbardi Caudell, Z. barberi
apterans. This fact has fueled the notion
that Zoraptera are cryptic, exceedingly rare,
4The Embiodea and Zoraptera together comprise clade
highly endemic, and occur in low abundances.
“Miroder Mystroptera Engel, new mirorder” as defined by
the characters established by Engel and Grimaldi (2000). Each of these assertions, however, is not
2006 RAFAEL AND ENGEL: NEW AMAZONIAN ZOROTYPUS 3
accurate. The apparent rarity of zorapterans SYSTEMATICS
stems in large part from the absence of
individuals seeking them or from a lack of Zorotypus (Zorotypus) amazonensis,
familiarity with the order when they are new species
stumbled across (Engel, 2004). Once a Figures 1-14
search image is developed for zorapterans,
Diagnosis: The new species is unique for
colonies can be located in almost any suitable
the combination of specialized, longitudinal,
habitat. Indeed, in the Amazon Basin zorap¬
paramedian rows of setae on the male vertex;
terans are rather common, and the Instituto
the clasperlike cerci of the male; the large
Nacional de Pesquisas da Amazonia now
female S8 with an apical cleft; the metafemur
has more than 1300 specimens collected
with 10 stiff, ventral spines and metatibia with
principally during the last three years. As
three ventral spines in both sexes; and hind
suggested by Engel (2001) for tropical species
wing with a distinct and true Cu vein (vide
and as demonstrated for Z. hubbardi by
Discussion, infra).
Hinojosa-Diaz et al. (in press), species can
Description: Apterous male (holotype).
have relatively large ranges, and appropriate
Body length (exclusive of antennae) 2.5 mm;
environmental and substrate conditions are
antennal length 1.8 mm. Integument light
readily available over these expansive distri¬
brown. Head subtriangular, posterior border
butions.
slightly broader than pronotum; compound
In the Neotropical region the Zoraptera
eyes absent, with a small black spot in position
previously consisted of merely 17 species. We
otherwise occupied by eyes; ocelli absent; head
herein describe a remarkable new species from
setae as depicted in figure 1; vertex with
the South American tropics, representing the
median area bordered by longitudinal, para¬
34th species for the order. The new species is
median rows of pronounced, dense, slender,
particularly noteworthy for the unusual de¬
and divergent setae (fig. 1); antenna (fig. 2)
velopment of the male cerci, which are
with first antennomere cup-shaped; combined
elongate, curved, and beset with three stiff
length of second and third antennomeres
setae along their inner margins making them
slightly shorter than that of first and fourth
appear clasperlike.
antennomeres; length of second antennomere
subequal to that of third antennomere; anten¬
MATERIALS AND METHODS nomeres IV-IX longer than wide, becoming
slightly shorter and paler distally. Labrum
Morphological terminology follows that of with setae in distal half (fig. 1). Mandible with
Engel (2000), with homologies for apical three short, pointed teeth and well-defined
abdominal structures based on unpublished molar region (fig. 1). Lacinia with apical comb
comparative data of M.S. Engel, while the (fig. 3). Maxillary palpus (fig. 4) five-segment¬
supraspecific classification follows that of ed, first palpomere (Pi) short, P2 slightly
Engel and Grimaldi (2000, 2002) and Engel longer than P3, P4 short and nearly one-half
(2003b) (table 1). The abbreviations P, T, and length P3, P5 longest, laterally compressed,
S are used for palpomere, tergum, and and twice as wide as P4, P5 nearly as long as
sternum, respectively. Specimens were macer¬ combined lengths of P3 and P4. Labial palpus
ated in hot 85% lactic acid and then dissected three-segmented, P2 shortest, one-third length
and drawn in glycerin in an excavated slide. of Pi and P3 individually, P3 laterally com¬
The entire body was mounted on slides in pressed, twice as wide as Pi.
balsam. In slide-mounted specimens the ab¬ Pronotum subrectangular, slightly broader
domen was removed and tergites and sternites than long, not constricted posteriorly; longest
were separated laterally and mounted with setae on lateral and anterior margins, smaller
their external surfaces facing upward. Some setae scattered on disc. Mesonotum slightly
sclerites were drawn after slide mounting. smaller than pronotum, with longest setae on
Type material is deposited in the Instituto lateral margins. Metanotum smaller then pre¬
Nacional de Pesquisas da Amazonia (INPA), ceding thoracic terga, distinctly wider than
Manaus, Brazil. long, with longest setae on lateral margins.
4 AMERICAN MUSEUM NOVITATES NO. 3528
Figs. 1-6. Male of Zorotypus amazonensis Rafael and Engel, new species. 1. Male head, dorsal view, with
. . . . .
antennae removed. 2 Antenna. 3 Lacinia. 4 Maxillary palpus. 5 Labial palpus. 6 Hind leg from
metafemur to metapretarsus. Scale bars = 0.1 mm except that those for figures 2 and 6 = 0.5 mm.
Legs with scattered setae of moderate length, thickened, rather oblique setae either side of
intermixed with several small setae. Pro- and midline and 4-5 typical setae more distally;
mesotibiae with one and two rather stout, T10 (figs. 7, 8) largely membranous, with
subapical setae ventrally, respectively. Hind small, median sclerite with pair of small setae,
leg (fig. 6) with metafemur about three times and lateral sclerites with 4-5 flanking, poste¬
longer than broad (as measured at its point of rior setae; epiproct nearly divided medially,
maximal breadth), slightly more swollen to¬ lateral portions somewhat triangular and
ward base than apex; posterior ventral surface extending around cereal base, medially strong¬
with 10 short, stiff spines, second spine about ly narrowed and fused to upcurved anal
twice length of others, distal spines smaller sclerite (mating hook) bearing anal tube;
and more closely spaced; metatibia with three paraprocts broadly attached to apicolateral
ventral, stiff spines evenly spaced in distal half. corners of epiproct (fig. 8). Cerci (fig. 7)
Abdominal T1 with single transverse row of unsegmented, elongate, clasperlike, with three
eight setae and two small, inconspicuous stout, pronounced setae on inner margin, first
lateral setae on each side of midline; T2-9 just apical of midpoint, second subapical,
with two transverse rows of 7-10 setae each third apical, all three setae directed inward;
side of midline; T9 (figs. 7, 8) with small other setae rather stout, long, and along outer,
median setae dorsolaterally followed by four lateral surface; basal half covered with short,
2006 RAFAEL AND ENGEL: NEW AMAZONIAN ZOROTYPUS 5
epiproct
paraproct
anal sclerite
& anal tube
S7
S8
11 12
Figs. 7-12. Zorotypus amazonensis Rafael and Engel, new species, abdominal structures. 7. Male
. . .
abdominal apex, dorsal view. 8 Male abdominal apex, apical view. 9 Male abdominal sterna. 10 Male
. .
sternum VIII and terminalia. 11 Female sternum VIII. 12 Female genital sclerite. Scale bars = 0.1 mm
except that for figure 9 = 0.5 mm. The terms paraproct and epiproct are used here to indicate the correct
sclerite homologies with other insects (based on unpublished comparative data of M.S. Engel) and in place
of the less formal terms employed by Bolivar y Pieltain and Coronado (1963). The anal sclerite is fused to
what we consider the epiproct in Zoraptera.
6 AMERICAN MUSEUM NOVITATES NO. 3528
dense spicules, such spicules absent at apex forewing length 2.7 mm, pterostigma present
and internally at base. Abdominal sterna as in as faint infuscation along anterior, apical wing
figure 9; first sternum not sclerotized and margin between R and Rs; R disappearing in
apparently absent; S2 divided into small, pterostigmal base (fig. 13); Rs nearly reaching
lateral sclerites, with two pairs of small setae to forewing apex, becoming faint and diffuse
on each side; S3-6 with two rows of short right near apex; rs-m present (fig. 13), very
setae, those apically and centrally slightly short, slightly shorter than first free abscissa
longer; S8 (hypandrium or subgenital plate) of Rs; M extending to posterior wing margin;
larger than preceding sterna (figs. 9, 10), CuAi present and extending to just beyond
rather oval, more sclerotized in apical portion, wing midpoint before terminating on posterior
distal margin produced medially, protuber¬ wing margin; CuA2 present as a short stub in
ance with two small, sub apical setae (fig. 10), basal third of wing; wing setae numerous on
setae scattered over disc and transverse row of membrane and margins, not clustered in any
longer setae at distal margin. region of wing; additional forewing details in
Terminalia (from paratype) as depicted in figure 13. Hind wing length 2.2 mm; venation
figure 10; terminalia principally membranous, typical, with stems of R and M starting
highly asymmetrical, and difficult to interpret separate near base and fusing for much of
except by two pieces: central, rather club- wing length, separating in distal third of wing,
shaped, more sclerotized portion and basal each vein terminating as evanescent vein on
portion that is less sclerotized and bearing an wing margin (fig. 14); short branch of Cu
apically directed, elongate, tubular projection present proximally and terminating as evanes¬
(the tubular portion may be the same structure cent vein on wing margin (fig. 14); exceedingly
that forms a dorsal, somewhat sclerotized and short and faint M+Cu stem present; additional
coiled tube in four neotropical species; i.e., Z. hind wing details in figure 14.
snyderi Caudell, Z. hamiltoni New, Z. mex- Holotype: Male (slide mounted), “BRASIL,
icanus Bolivar y Pieltain, and Z. cramptoni AM [Amazonas], Itacoatiara, Madeireira Mil,
Gurney). 17.viii.2002, J.A. Rafael & J. Vidal, sob casca
Apterous female. As described for the male de tronco caido” (INPA).
except as follows: body length (exclusive of Paratypes: One male, three females (two
antennae) 2.7 mm; antennal length 2.0 mm. slide mounted, one in alcohol), same data
Head without central, paramedian row of as holotype (INPA). One alate female
pronounced cluster of divergent, slender setae (slide mounted), “Manaus, Reserva Ducke,
on vertex. Metafemur ornamentation slightly 21 .ii. 1992, Igarape Barro Branco, arm. adesi-
weaker than that of male. Abdominal T9-10 va, J. Vidal” (INPA). One alate female (in
normal, without modifications; S8 larger than alcohol), “Manaus, ZF-3, km-23, Fazenda
preceding sterna, slightly asymmetrical, more Esteio, 18.iii. 1987, B.C. Klein, armadilha
sclerotized in distal third, with large distal, malaise” (INPA).
median cleft (fig. 11). Cerci normal, small, Etymology: The specific epithet refers to
slightly curved proximally, with rather stout the region in which specimens were collected,
setae but no spiniform setae as in male. i.e., Amazonia.
Genital fork small, slightly sclerotized Comments: Among Brazilian species, Z.
(fig. 12). amazonensis is among the most completely
Alate female. As described for the male characterized (table 2), with both sexes and
except as follows: body dark brown to black; morphs documented. Within this fauna, Z.
compound eyes present and black; three ocelli amazonensis is apparently most similar to Z.
present on vertex; venation generally reduced shannoni Gurney, also described from Brazil,
and weak, most veins nebulous (i.e., mostly in that both have modified and slender setae
represented by distinctly fuscous lines on on the vertex. The organization of these setae
otherwise hyaline or lightly infuscate mem¬ differs prominently (vide Description, supra,
brane) although veins R, Rs, and M in and Discussion, infra), serving to separate the
forewing and R+M in hind wing at least two species readily in the male sex. Both
weakly tubular for portions of their length; species also have similar metafemoral orna-
2006 RAFAEL AND ENGEL: NEW AMAZONIAN ZOROTYPUS 7
.
Figs. 13-14. Female alate of Zorotypus amazonensis Rafael and Engel, new species. 13 Forewing
.
(2.7 mm in length). 14 Hind wing (2.2. mm in length).
mentation, although with 10 stiff, ventral are unknown, hindering a comprehensive
spines in Z. amazonensis and six such spines study of particular character systems.
in Z. shannoni. Likewise, both species have Appendix 1 summarizes the known forms of
a membranous endophallus. the world’s species. Despite this impediment,
some preliminary conclusions can be drawn
DISCUSSION (e.g., Engel and Grimaldi, 2000; Engel,
2003b), and the discovery of Z. amazonensis
Owing to the paucity of studies on permits some new interpretations of anatom¬
Zoraptera and the dimorphic nature of the ical traits in Zorotypus. It is fortunate that not
species, comparisons across species are chal¬ only males and females are known for Z.
lenging. Species may not only be known only amazonensis but also apterous and alate forms
from a single sex, but sometimes alate forms have been recovered, permitting a more com-
TABLE 2
Checklist of Brazilian Zoraptera (updated from Rafael and Engel, in press)
Species Sexes known Alate known Nymph known Geographical records®
Z. amazonensis Q'Q X X Brazil (AM)
Z. brasiliensis 9 X X Brazil (MG, RJ, SP, PR, SC)
Z. huxleyi a? — — Peru, Brazil (AM)
Z. shannoni cr X — Brazil (MT, AM)
Z. weidneri erg — — Brazil (AM)
aStates of Brazil: AM = Amazonas, MG = Minas Gerais, MT = Mato Grosso, RJ = Rio de Janeiro, SP = Sao Paulo,
PR = Parana, and SC = Santa Catarina.
AMERICAN MUSEUM NOVITATES NO. 3528
plete characterization of this species as well as phylogenetic context. For example, both Z.
a more meaningful comparison with taxa that delamarei and Z. congensis Ryn-Tournel have
are similarly documented. a clearly defined fontanelle and, aside from the
New (1978) noted that the cluster of setae presence of such an obvious orifice (e.g., vide
on the vertex of Z. shannoni is unusual and Delamare-Deboutteville, 1951), they may be
suggested that these setae were homologous further united by the diffuse, sinuous setae
with those present on Z. delamarei Paulian anterior to the fontanelle. In Z. barberi and Z.
from Madagascar. The homology of these two shannoni, however, there is a different condi¬
features, however, appears dubious given the tion. The fontanelle is not clearly demarked as
available data (Engel, personal obs.). In Z. a distinct opening; instead, in each of these
shannoni the male vertex lacks a fontanelle but species it can be recognized as a tight cluster of
has a distinct, dense, circular cluster of long, modified setae on the vertex, sometimes set
sinuous setae standing out among the usual into a faint depressed region and clearly with
distribution of vertical setae that are slightly porous integument (e.g., vide New, 1978;
shorter, straight (although not erect, frequent¬ Choe, 1989). This overall form of glandular
ly suberect in position), and more rigid. The opening may be homologous between the two
vertex of male Z. delamarei, by constrast, has species, with the particular structure of the
a distinct fontanelle orifice that is anteriorly individual setae being distinctive to each (long,
bordered by a loose collection of short, sinuous in Z. shannoni', shorter, stout, and
sinuous setae. These setae are not congregated erect in Z. barberi). Under this scenario, the
into a distinct patch and are generally the presence of the gland and the glandular
same size or shorter than the surrounding opening may be homologous across species,
typical vertical setae. Indeed, the setal patch of while the particular form of the opening and
Z. shannoni likely serves a glandular function surrounding setae may only be homologous
during courtship in which males offer secre¬ for subsets of such taxa. This is not to say at
tions from a vertical gland to the female prior this time that Z. barberi and Z. shannoni are
to copulation (as of yet, this has not been sister taxa (indeed the species are quite
conclusively demonstrated but appears as¬ different) since the distribution of vertical
suredly correct based on examination of the structures across other Zorotypus is poorly
structure), like that documented for Z. barberi documented and ultimately may alter some
(Choe, 1995). Such is assuredly the case in Z. of the preliminary ideas developed here.
delamarei and other species bearing a more However, among those few species in which
distinct fontanelle orifice. While the gland the male vertex has been examined, the
(where observed), its secretions, and certainly structures seen in these two neotropical species
the presence of a fontanelle are likely homol¬ more closely approximate each other than
ogous, the homology of specialized setae on either compare to the condition of the African
the vertex and surrounding and glandular species. Certainly a more extensive compara¬
openings (either as a well-defined fontanelle tive study of fontanelle structures is required
orifice or as small region of porous integument within a phylogenetic framework before de¬
and clustered setae) is less obvious. It may be finitive conclusions can be reached. Despite
that numerous, species-specific modifications this current hindrance, these preliminary ob¬
have arisen in order to channel or capture servations are tantalizing and indicate that
droplets of secretions emerging from a vertical numerous modifications await more definitive
gland as offerings to females during courtship. study and are likely associated with interesting
Alternatively, these setae may indeed prove to aspects of zorapteran mating behavior.
be homologous but may assume various forms
in different species. It seems more likely,
5Note that the presence of a vertical gland has not yet
however, that different setal configurations been conclusively documented across the entire order,
may unite sets of species5 within Zorotypus s. although it would seem that it is a feature of all Zorotypus.
Alternatively, it may be that a monophyletic subset of
str., and that while such features may be
Zorotypus has the vertical gland and that within this clade
homologous among subsets of species, these
a diversity of forms of orifices and surrounding setae has
types may prove to be independent in a larger arisen.
2006 RAFAEL AND ENGEL: NEW AMAZONIAN ZOROTYPUS 9
The vertex of Z. amazonensis also bears male of Z. amazonensis is a remarkable
a cluster of modified setae and, like the other autapomorphy for this species and one in
American species studied to date, it lacks the which the study of the biology and functional
obvious fontanelle orifice of Z. delamarei and morphology would be of great significance.
Z. congensis. However, despite the overall The new species also highlights the need for
similarity of this species to Z. shannoni, the extensive exploration of the tropical zones
organization of these setae in Z. amazonensis throughout the world for additional material
is quite distinctive and utterly unique within and species of Zoraptera. While much work
the order. Rather than being clustered into remains within the neotropical region, such
a tight, circular patch as in Z. shannoni or Z. fieldwork is even more intensely needed in
barberi, the setae are arranged into tight, Africa and Asia where only few, scattered
longitudinal, paramedian rows on the vertex records exist across vast regions with suitable,
if not ideal, zorapteran habitat. The potential
(fig. 1).
Aside from an unusual cephalic arrange¬ for new species and thereby new insights into
this poorly understood order is great. Indeed,
ment of setae, Z. amazonensis is further unique
it is possible that with adequate sampling the
among those species for which alates are
diversity of the order may eventually more
known by having a distinct branch of Cu
than double.
proximally in the hind wing (fig. 14). The
recognition of this vein permits the determi¬
nation that the hind wing posterior stem ACKNOWLEDGMENTS
termed when present as “Cu” by previous
authors (e.g., Choe, 1989; Engel and Grimaldi, We are thankful for the efforts of two
2000, 2002) is, in fact, vein M. This partially anonymous reviewers who carefully studied
supports the conclusion of Kukalova-Peck the manuscript prior to publication. The
and Peck (1993) that this vein is part of the senior author (a fellow of CNPq) is grateful
medial system, but we cannot agree that it to Joao Ferreira Vidal and Francisco Felipe
represents MP, and their more elongate, Xavier Filho, both from INPA, for support
hypothetical stem “MP+Cu” extending to and assistance in collecting zorapterans, and
to CNPq (process number 472978/03-9) for
the extreme wing base (e.g., their fig. 6) is
partially financing the collections. The junior
not supported by observations of this or other
author is grateful to Ismael A. Hinojosa-Diaz
species (Engel, personal obs.). There is also
for assistance during portions of this work.
a short stub proximal to the separation of the
This is contribution No. 3427 of the Division
true cubital vein, indicating that M and Cu are
of Entomology, University of Kansas Natural
fused in the hind wing base. While most
History Museum.
species have the short basal stem of M present
(again, termed Cu by previous authors)
(fig. 14), the presence of a short, longitudinal REFERENCES
vein posterior to the main vein R+M is unique
Arillo, A., and M.S. Engel. In press. Rock crawlers
among species with documented venation.
in Baltic amber (Notoptera: Mantophas-
This vein branches very early in the wing
matodea). American Museum Novitates.
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Beutel, R.G., and S.N. Gorb. 2001. Ultra structure
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(fig. 14). The discovery of the alate of Z. (Arthropoda): evolutionary patterns inferred
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plete and accurate homologization of hind Zoological Systematics and Evolutionary
wing venation in Zoraptera. Research 39(4): 177-207.
Bolivar y Pieltain, C., and L. Coronado G. 1963.
The discovery of Z. amazonensis demon¬
Estudio de un nuevo Zorotypus proveniente de
strates that significant discoveries remain to be
la region Amazonica Peruana (Ins., Zoraptera).
made in the order Zoraptera and that as-of-yet
Ciencia (Mexico ity) 22(4): 93-100.
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Choe, J.C. 1989. Zorotypus gurneyi, new species,
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