ArticlePDF Available

Taxonomic revision of the myrmecophilous, meliponiphilous and rhizophilous soft scale genus Cryptostigma Ferris (Hemiptera: Coccoidea: Coccidae)

Authors:

Abstract

The present study revises the soft scale insects of the genus Cryptostigma Ferris (Hemiptera: Coccoidea: Coccidae), which comprises a group of New World species associated with ants or bees or of hypogeal habit. It includes 17 species, of which 10 species are new to science. The study is based on the external, mostly cuticular morphology of the adult females and first-instar nymphs. Taxonomic keys based on adult females and known first-instar nymphs are provided. For each species a description or redescription is given, followed by information on its distribution, host plants, and known biological information including associated ants or bees. The new species described are Cryptostigma chacoensis sp. nov., C. gullanae sp. nov., C. jonmartini sp. nov., C. longinoi sp. nov., C. melissophilum sp. nov., C. mexicanum sp. nov., C. philwardi , sp. nov. C. rhizophilum sp. nov., C. serratum sp. nov., and C. tuberculosum sp. nov. Neolecanium urichi (Cockerell) is transferred to Cryptostigma as C. urichi (Cockerell) comb. nov., and C. quinquepori (Newstead) is synonymized with Cryptostigma urichi, comb. nov. Lectotypes are designated for Cryptostigma saundersi Laing, Lecanium silveirai Hempel, Lecanium urichi Cockerell and Akermes quinquepori Newstead. Cryptostigma inquilina (Newstead) is amended to C. inquilinum in order to match the neutral ending “stigma”. The following names are treated as nomina nuda: Cryptostigma jamaicensis and Lecanopsis jamaicensis (Ben-Dov, 1993: 97). Key words: Cryptostigma, taxonomic keys, Myzolecaniinae, soft scales
Accepted by C. Hodgson: 20 Sept 2010; published: 3 Dec. 2010
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334 (online edition)
Copyright © 2010 · Magnolia Press
Zootaxa 2709: 172 (2010)
www.mapress.com
/zootaxa/
Monograph
ZOOTAXA
Taxonomic revision of the myrmecophilous, meliponiphilous
and rhizophilous soft scale genus Cryptostigma Ferris
(Hemiptera: Coccoidea: Coccidae)
TAKUMASA KONDO
Corporación Colombiana de Investigación Agropecuaria (CORPOICA), Centro de Investigación Palmira, Calle 23, Carrera 37,
Continúo al Penal, Palmira, Valle, Colombia. E-mail: tkondo@corpoica.org.co
Magnolia Press
Auckland, New Zealand
2709
KONDO
2 · Zootaxa 2709 © 2010 Magnolia Press
Takumasa Kondo
Taxonomic revision of the myrmecophilous, meliponiphilous and rhizophilous soft scale genus Cryp-
tostigma Ferris (Hemiptera: Coccoidea: Coccidae)
(Zootaxa 2709)
72 pp.; 30 cm.
3 December 2010
ISBN 978-1-86977-637-4 (paperback)
ISBN 978-1-86977-638-1 (Online edition)
F
IRST PUBLISHED IN 2010 BY
Magnolia Press
P.O. Box 41-383
Auckland 1346
New Zealand
e-mail: zootaxa@mapress.com
http://www.mapress.com/zootaxa/
© 2010 Magnolia Press
All rights reserved.
No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any
means, without prior written permission from the publisher, to whom all requests to reproduce copyright
material should be directed in writing.
This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose
other than private research use.
ISSN 1175-5326 (Print edition)
ISSN 1175-5334 (Online edition)
Zootaxa 2709 © 2010 Magnolia Press · 3
REVISION OF CRYPTOSTIGMA FERRIS
Table of contents
Abstract ...............................................................................................................................................................................3
Resumen ..............................................................................................................................................................................3
Introduction......................................................................................................................................................................... 4
Association of Cryptostigma species with ants, stingless bees and plant root systems ......................................................5
Material and methods ..........................................................................................................................................................5
Key to separate Cryptostigma Ferris from morphologically similar Houardia Marchal and Myzolecanium Beccari based
on adult female and first-instar nymphs ...................................................................................................................... 7
Key to separate the male and female instars of Cryptostigma Ferris.................................................................................. 7
Cryptostigma Ferris .............................................................................................................................................................8
Key to the adult females of Cryptostigma Ferris ............................................................................................................... 9
Key to the known first-instar nymphs of Cryptostigma Ferris .........................................................................................10
Description of species .......................................................................................................................................................11
Cryptostigma biorbiculus Morrison ...........................................................................................................................11
Cryptostigma chacoensis Kondo, sp. nov. ................................................................................................................15
Cryptostigma guadua Kondo & Gullan .................................................................................................................... 19
Cryptostigma gullanae Kondo, sp. nov. ....................................................................................................................23
Cryptostigma inquilinum (Newstead)........................................................................................................................ 25
Cryptostigma jonmartini Kondo, sp. nov. ................................................................................................................ 30
Cryptostigma longinoi Kondo, sp. nov. .....................................................................................................................34
Cryptostigma melissophilum Kondo, sp. nov. ..........................................................................................................37
Cryptostigma mexicanum Kondo, sp. nov. ................................................................................................................41
Cryptostigma philwardi Kondo, sp. nov. ..................................................................................................................43
Cryptostigma reticulolaminae Morrison ................................................................................................................... 45
Cryptostigma rhizophilum Kondo, sp. nov.
..............................................................................................................49
Cryptostigma saundersi Laing ...................................................................................................................................54
Cryptostigma serratum Kondo, sp. nov. ...................................................................................................................56
Cryptostigma silveirai (Hempel) ..............................................................................................................................58
Cryptostigma tuberculosum Kondo, sp. nov. ............................................................................................................62
Cryptostigma urichi (Cockerell) comb. nov. .............................................................................................................66
Acknowledgements ...........................................................................................................................................................70
References .........................................................................................................................................................................71
Abstract
The present study revises the soft scale insects of the genus Cryptostigma Ferris (Hemiptera: Coccoidea: Coccidae),
which comprises a group of New World species associated with ants or bees or of hypogeal habit. It includes 17 species,
of which 10 species are new to science. The study is based on the external, mostly cuticular morphology of the adult
females and first-instar nymphs. Taxonomic keys based on adult females and known first-instar nymphs are provided.
For each species a description or redescription is given, followed by information on its distribution, host plants, and
known biological information including associated ants or bees. The new species described are Cryptostigma chacoensis
sp. nov., C. gullanae sp. nov., C. jonmartini sp. nov., C. longinoi sp. nov., C. melissophilum sp. nov., C. mexicanum sp.
nov., C. philwardi , sp. nov. C. rhizophilum sp. nov., C. serratum sp. nov., and C. tuberculosum sp. nov. Neolecanium
urichi (Cockerell) is transferred to Cryptostigma as C. urichi (Cockerell) comb. nov., and C. quinquepori (Newstead) is
synonymized with Cryptostigma urichi, comb. nov. Lectotypes are designated for Cryptostigma saundersi Laing,
Lecanium silveirai Hempel, Lecanium urichi Cockerell and Akermes quinquepori Newstead. Cryptostigma inquilina
(Newstead) is amended to C. inquilinum in order to match the neutral ending “stigma”. The following names are treated
as nomina nuda: Cryptostigma jamaicensis and Lecanopsis jamaicensis (Ben-Dov, 1993: 97).
Key words: Cryptostigma, taxonomic keys, Myzolecaniinae, soft scales
Resumen
El presente estudio revisa las escamas blandas del género Cryptostigma Ferris (Hemiptera: Coccoidea: Coccidae), cual
está compuesto por especies del Nuevo Mundo asociadas con hormigas, abejas sin aguijón o de hábitos subterraneos. El
KONDO
4 · Zootaxa 2709 © 2010 Magnolia Press
género incluye 17 especies de las cuales 10 son nuevas para la ciencia. El presente estudio se basa principalmente en la
morfología cuticular externa de las hembras adultas y ninfas del primer instar. Se proveen claves taxonómicas para las
hembras adultas y las ninfas del primer instar. Para cada especie se hace una descripción o redescripción, con
información sobre su distribucion geográfica, plantas hospederas, y lo que se conoce sobre su biologia, incluyendo sus
asociasiones con hormigas y abejas sin aguijón. Las nuevas especies descritas en este estudio son: Cryptostigma
chacoensis sp. nov., C. gullanae sp. nov., C. jonmartini sp. nov., C. longinoi sp. nov., C. melissophilum sp. nov., C.
mexicanum sp. nov., C. philwardi , sp. nov. C. rhizophilum sp. nov. , C. serratum sp. nov., y C. tuberculosum sp. nov.
Neolecanium urichi (Cockerell) es transferida a Cryptostigma como C. urichi (Cockerell) comb. nov., y C. quinquepori
(Newstead) se sinonimiza con Cryptostigma urichi, comb. nov. Se designan lectotipos para Cryptostigma saundersi
Laing, Lecanium silveirai Hempel, Lecanium urichi Cockerell y Akermes quinquepori Newstead. Cryptostigma
inquilina (Newstead) se emenda a C. inquilinum para hacer concordar con la terminación neutral “stigma”. Los
siguientes nombres se tratan como nomina nuda: Cryptostigma jamaicensis y Lecanopsis jamaicensis Ben-Dov, 1993:
97.
Introduction
The scale insects include approximately 7,355 species distributed in 1,050 genera and 28 (22 extant + 6 fossil)
families (Ben-Dov et al., 2010). The Coccidae or soft scales make up the third most species rich family with
approximately 1,113 species in 163 genera (Ben-Dov et al., 2010). Scale insects occur anywhere where plants
can grow, with species occurring even in the cold arctic tundra. On the host, they are found on the leaves,
branches, trunk, under the bark, and on the fruits and roots. Some species may cause various kinds of plant
deformities including chlorotic spots, pits and galls (Ben-Dov et al., 2010).
The genus Cryptostigma is currently included in the subfamily Myzolecaniinae, a group erected by
Hodgson (1994) to include species with the following combination of features: lack of (i) dorsal tubular ducts
and (ii) eyespots; and the presence of: (iii) anal plates with numerous setae on dorsal surface, (iv) particularly
large spiracles, with broad bands of spiracular disc-pores between margin and spiracles, (v) ventral tubular
ducts of only one type, frequently restricted to a group on each side of genital opening, (vi) bands of (often
rather spinose) setae replacing the normal pairs of long pregenital setae, (vii) reduced legs with claw digitules,
(viii) reduced antennae, and (ix) a short anal tube. According to Hodgson (1994), members of the
Myzolecaniinae are present in all zoogeographical regions, except China, the Palearctic Region, and New
Zealand. The Myzolecaniinae originally included 16 genera: Akermes, Alecanium, Alecanopsis,
Cribrolecanium, Cryptostigma, Cyclolecanium, Halococcus, Houardia, Megasaissetia, Myzolecanium,
Neolecanium, Paractenochiton, Pseudophilippia, Richardiella, Toumeyella and Xenolecanium (Hodgson,
1994). Later, four new genera: Aztecalecanium Kondo & Williams (2004), Octolecanium Kondo (Kondo et
al., 2005a), Neotoumeyella Kondo & Williams (2009) and Torarchus Gullan & Stewart (1996) were added.
Kondo et al. (2005b) pointed out that some species in the Paralecaniini (Coccinae) shared many features with
those in the Myzolecaniinae, and transferred Xenolecanium to the subfamily Coccinae, based on
synapomorphic features of the adult females and first-instar nymphs with members of the Paralecaniini,
bringing the total number of genera in the Myzolecaniinae to 19.
Cryptostigma was revised previously by Qin and Gullan (1989), who recognized 7 species including 2
new Australian species, C. endoeucalyptus and C. magnetinsulae. Gullan et al. (1993) later transferred the 2
species described by Qin and Gullan (1989), and a third species, C. robertsi Williams & Watson (1990)
described from Papua New Guinea, to the rediscovered genus Myzolecanium Beccari. Gullan et al. (1993)
also suggested that Cryptostigma could become a junior synonym of the much older name Myzolecanium, but
that a systematic revision of the group would be necessary before making such a decision. Hodgson (1994)
also considered Cryptostigma to be close to Myzolecanium, based on comparison of the type species of each
genus. He pointed out that Cryptostigma lacks cribriform plates, and the pregenital disc-pores are restricted to
a small area around the vulva in Myzolecanium and the ventral microducts have swollen ductules. Here
Cryptostigma is considered as a separate genus from Myzolecanium, and a key to separate Cryptostigma from
the morphologically similar Houardia Marchal and Myzolecanium is provided. The present study has included
all currently known species of Cryptostigma but, because of the secluded habit of these insects (e.g., found
inside hollow branches and underground), many more species are expected to be found in nature.
Zootaxa 2709 © 2010 Magnolia Press · 5
REVISION OF CRYPTOSTIGMA FERRIS
Association of Cryptostigma species with ants, stingless bees and plant root systems
Most members of the genus Cryptostigma are found internally on their host plants in hollow branches, or
trunks in association with ants (Newstead 1917, 1920; Ferris, 1922; Morrison, 1922, 1929; Laing, 1925) and
rarely stingless bees (Camargo and Pedro, 2002a, b). According to Qin and Gullan (1989), Cryptostigma is
involved with four ant subfamilies: Dolichoderinae, Formicinae, Myrmicinae and Pseudomyrmecinae.
Although Qin and Gullan’s (1989) revision of Cryptostigma included members of the genus Myzolecanium
(i.e., M. endoeucalyptus (Qin & Gullan) and M. magnetinsulae (Qin & Gullan), then included in
Cryptostigma), these same four ant subfamilies were found to be associated with Cryptostigma sensu stricto.
Myzolecanium endoeucalyptus is tended by ants, probably Crematogaster sp. (Myrmicinae) or Iridomyrmex
sp. (Dolichoderinae), and M. magnetinsulae has no ant association records (Qin & Gullan, 1989). Species
known to be associated with Cryptostigma are: Azteca alfari cecropiae (Morrison, 1929); Azteca longiceps
(Morrison, 1929); Camponotus sp., C. longipilis, C. mirabilis (Kondo & Gullan, 2004), C. novogrenadensis
(Morrison, 1922); Crematogaster brevispinosa (Newstead, 1920); Myrmelachista sp. (Morrison, 1929),
Myrmelachista ambigua ramulorum Wheeler (Ferris, 1922); Pseudomyrmex sericea (as Pseudomyrma
sericea) (Morrison, 1929), and P. viduus (Ward, 1999). Other ants found associated with Cryptostigma are:
Azteca spp., Azteca nigricans, A. xanthochroa, Crematogaster stolli, Pseudomyrmex spp., Pseudomyrmex sp.
cf. fortis (unpublished data). Two of the new species are found closely associated with stingless bees (Apidae:
Meliponini). One, C. melissophilum sp. nov., is tended by the stingless bee Schwarzula coccidophila
(Camargo & Pedro, 2002b) and the other species, C. chacoensis sp. nov. is found inside the nests of two
stingless bees, Schwarzula timida and Plebeia sp. (David Roubik personal communication). Two other
species: C. silveirai, comb. nov. and C. rhizophilim
sp. nov. are hypogeal in habit and are not known to be
tended by hymenopterans.
Material and methods
Dry and slide-mounted material was borrowed from various institutions. Specimens were slide mounted using
methods discussed by Kosztarab (1996) and Williams and Granara de Willink (1992). Measurements were
made using an ocular micrometer on an Olympus (JAPAN) BX40 phase-contrast microscope. Descriptions
are based on multiple slide-mounted specimens. Total body length is presented in millimeters whereas all
other measurements are presented in micrometers (
m) as a range. Scale insect illustrations show the dorsal
surface on the left half and the ventral surface on the right half. Important details are enlarged and placed
around the perimeter of the main figure. The dermal structures and enlargements are not in direct proportion
to each other. Listed for each species treatment under “Material studied” are the locality, collection date,
collector, host and other biological information, code numbers when present, and number of slides and
specimens. The first number indicates the number of slides and the second number (in parentheses) the
number of specimens. Also noted are the type of material, (i.e., primary type, paratypes or others), and the
depository abbreviated in parentheses.
The terminology used to describe the adult female follows mostly that of Hodgson (1994) and for the
first-instar nymph that of Kondo et al. (2005). The following terms are introduced here to describe
morphological features commonly seen in Cryptostigma.
Cribriform platelets: Very small cribriform plates are herein termed cribriform platelets. Each cribriform
platelet is generally less than 20
m at widest point, and can be found singly or in groups.
Orbicular pores: “Simple orbicular pores” are dorsal circular structures composed of a thin membrane
often surrounded by a sclerotized rim. “Compound orbicular pores” are composed of a thin membrane
associated with small simple pores, bilocular microducts and slender setae. Orbicular pores can be found
singly on the dorsum or incorporated into the spiracular sclerotization. The function of the orbicular pores is
unknown, although they may serve as sensory and/or secretory organs involved in the communication
between the scale insect and the mutualistic hymenopteran. Qin and Gullan (1989) used the name “compound
pores” to refer to these pores. The name “orbicular pore” coined here is derived from the species epithet of C.
KONDO
6 · Zootaxa 2709 © 2010 Magnolia Press
biorbiculus Morrison, a species with two orbicular pores. Orbicular pores should not be confused with
cribriform plates, which are sclerotized plates that have numerous similar pores in them (Hodgson, 1994).
Sclerotic pores: These are pores surrounded by sclerotized areas of various shapes, usually circular or
rectangular, found on the dorsal derm.
Stigmatic sclerotization: An area of sclerotization, which is closely associated with the spiracular
openings.
Submarginal setae: These generally refer to the ventral setae located along the submargins of the insect
body. In first-instar nymphs, these are usually found in pairs in the thorax and abdomen; each pair of
submarginal setae is composed of an inner and outer submarginal seta. The submarginal setae furthest from
the margin are the inner submarginal setae and those closest to the margin are called outer submarginal setae.
Abbreviations used in the figures are as follows: Adult female: anal plate (aplt); anal ring (ar); antenna
(ant); apodeme (apd); cribriform platelets (crbpltlts); dorsal microduct (dm); dorsal setae (dset); enlargement
of margin (em); large dorsal microduct (ldm); enlargement of dorsal derm (edd); enlargement of marginal
derm (emd); large simple pore (lsp); marginal seta (mset); small dorsal microduct (sdm); small simple pore
(ssp); orbicular pore (orbp); pregenital disc-pore (pdp); preopercular pore (prp); simple pore (sp); sclerotic
pore (sclrp); spiracular disc-pore (spdp); stigmatic sclerotization (stgsclr); stigmatic spines (stgsp); ventral
microduct (vm); ventral tubular duct (vtbd); ventral seta (vset). First-instar nymph: anal plate (aplt); anal ring
(ar); dorsal microduct (dm); dorsal seta (dset); marginal seta (mset); simple pore (sp); spiracular disc-pore
(spdp); stigmatic spines (stgsp); submarginal setae (sbset); trilocular pore (trp); ventral microduct (vm);
ventral cephalic seta (vcset); ventral outer submarginal seta (vosset).
Specimen depositories
AUCC Auburn University Coccoidea Collection, Auburn University, Auburn, Alabama, USA
BME Bohart Museum of Entomology, University of California, Davis, USA
BMNH Natural History Museum, London, England
CSCA California State Collection of Arthropods at California Department of Food and Agriculture,
Sacramento, USA.
IBSP Instituto Biologico de São Paulo, São Paulo, Brazil
MNHN Museum National d’Histoire Naturelle, Paris, France
MUSM Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru
MZSP Museu de Zoologia, Universidade de São Paulo, São Paulo, Brazil
NZAC New Zealand Arthropod Collection, Landcare Research, Auckland, New Zealand
RPSP Ribeirão Preto, Universidade de São Paulo, São Paulo, Brazil
SANC South African National Collection of Insects, Pretoria, South Africa
UNAD Museo de Entomologia, Universidad Nacional Agraria, Apartado 456, La Molina, Lima, Peru
USNM The National Museum of Natural History Coccoidea Collection, Beltsville, Maryland, USA
Material of species of other genera studied for comparison and construction of keys
Houardia abdita De Lotto. Adult female, South Africa, Pretoria, Transvaal, 12.xi.1963, coll. J. Munting,
ex Burkea africana, H.C. No. 763, 5,7,11,15, 4(4), Paratypes (SANC).
Houardia mozambiquensis Hodgson. Mozambique, Barra Falsa, 25.xi.1965, coll. not given, ex Casuarina
cunninghamiana, No. 10a, C.I.E. 9192, A1257, B.M. 1968. 27, 1(5: 1 adult female + 4 first-instar nymphs),
Holotype and Paratypes (BMNH).
Houardia troglodytes Marchal. Zimbabwe, Salisburg, Gilston Farm, 1949, coll. not given, ex inside
tunnels of Burkea africana, det. C.J. Hodgson, 1989, 2(2), (BMNH); Ivory Coast, Savane de Lanto, viii.1974,
coll. B. Darchen, ex Conopterix?, 5(8: 4 adult females + 4 first-instar nymphs), No. 6116-2, (MNHN); Sudan,
G.R.S., Medani, 21.xi.1961, coll. L.R. Schultz, det. D.J. Williams, ex Ficus, No. 4641-18069, B.M. 1963-3,
1(4 first-instar nymphs) (BMNH).
Zootaxa 2709 © 2010 Magnolia Press · 7
REVISION OF CRYPTOSTIGMA FERRIS
Myzolecanium endoeucalyptus (Qin & Gullan). Australia, New South Wales, Monga, 27.x.1986, coll.
Milton Lewis & Andrew Cockburn, ex on hollow tree branch of Eucalyptus viminalis, 4(4: 2 adult females +
2 first-instar nymphs), Paratypes (NZAC); Australia, Canberra, ANBG, near Section 77, 15.x.1987, coll. S.
Donaldson, 2(2: 1 adult female + 1 first-instar nymph), Paratype (NZAC).
Myzolecanium magnetinsulae (Qin & Gullan). Australia, Queensland, Magnetic Island, 12.xi.1922, coll.
G. F. Hill, # 1587, host not given, 1(1) (BMNH). Australia, Queensland, Magnetic Island, “1088/WWF” &
“Magnetic Isd/Queensland/G.F. Hill/ 1587 /12.xi.1922, det. P.J. Gullan, 1999, 1(1 first-instar nymph)
(BMNH).
Myzolecanium robertsi (Williams & Watson). Papua New Guinea, Madang Province, Baku, G.R.C.
Plantations, 8.viii.1984, coll. H. Roberts, ex on trunk of Terminalia brasii, (D.P.I) No. 2576, C.I.E. A. 16262,
2(4: 2 adult females + 2 first-instar nymphs), Paratypes (BMNH).
Key to separate Cryptostigma Ferris from morphologically similar Houardia Marchal and Myzoleca-
nium Beccari based on adult female and first-instar nymphs
1. Adult female. Pregenital disc-pores completely absent. With several subcircular groups of simple disc-pores present
submarginally. Anal plates together pyriform. First-instar nymph. Marginal setae strongly bent near apex. Stigmatic
setae present in two opposite groups on each edge of a deep stigmatic cleft. Anal plates together pyriform. Known
from the Ethiopian region................................................................................................................. Houardia Marchal
- Adult female. Pregenital disc-pores present. Without several subcircular groups of simple disc- pores present sub-
marginally, although a single subcircular group of pores may occur laterad to each spiracular sclerotization. Anal
plates together quadrate or subcircular, rarely pyriform. First-instar nymph. Marginal setae usually straight, rarely
with strongly bent apex. Stigmatic setae present on deepest part of stigmatic cleft, not present in two opposite groups
on each edge of stigmatic cleft. Anal plates together quadrate. Not known from the Ethiopian region......................2
2. Adult female. Dorsal microducts with very long inner ductules (about 20 times the length of the width of the outer
rim of the duct), except for M. kibarae in which they are short and similar to ventral microducts; pregenital disc-
pores always restricted to small area around vulva; stigmatic setae present or absent; if present, numerous (20–100);
ventral microducts appearing bilocular, with long inner ductules; legs either reduced or completely absent; anal
plates with 15–50 setae on dorsal surface. First-instar nymph. Antennae 6 segmented; with a seta present near each
coxa; area just above anal plates membranous, not showing signs of sclerotization; with 6 ventral submedian setae
(known from the Australasian region)........................................................................................Myzolecanium Beccari
- Adult female. Dorsal microducts not as long as above, usually each ductule less than 10 times the length of the width
of the outer rim of the duct. Pregenital disc-pores either restricted to around vulva or extending onto anterior abdom-
inal segments; stigmatic setae either absent or less than 3; ventral microducts appearing unilocular, with short inner
ductules (except for C. reticulolaminae Morrison in which they are as long as its dorsal microducts); legs reduced,
often represented by a group of setae and a small sclerotic plate, never completely absent, although often hard to
detect; anal plates with 4–21 setae on dorsal surface. First-instar nymph. Antennae 5 or 6 segmented; without a seta
present near each coxa; with 1 or 2 membranous folds just above anal plates; usually with 3 ventral submedian setae,
but rarely up to 5, or many setae present on all abdominal segments and thorax, never in 6 pairs (known from the
Neotropical region).........................................................................................................................Cryptostigma Ferris
Key to separate the male and female instars of Cryptostigma Ferris
1. Insects with wings or wing buds..........................................................................Adult male, or male pupa or prepupa
- Insects without wings or wing buds..............................................................................................................................2
2. Legs and antennae long, well developed, showing no reduction of segments; legs clearly cursorial..........................3
- Legs and antennae short, greatly reduced; legs not cursorial.......................................................................................5
3. Each anal plate with a long apical seta, if apical seta broken, large setal socket of apical seta present on apex of each
plate.................................................................................................................................................... first-instar nymph
- Each anal plate with neither a long apical seta nor with a large setal socket on apex of each plate ............................4
4. Dorsum with tubular ducts....................................................................................................second-instar nymph male
- Dorsum without tubular ducts............................................................................................second-instar nymph female
KONDO
8 · Zootaxa 2709 © 2010 Magnolia Press
5. Vulva and multilocular disc-pores absent around vulva ..................................................................third-instar nymph
- Vulva present; multilocular disc-pores present in perivulvar region..........................................................adult female
Notes. No males are known from Houardia and Myzolecanium, and these two genera appear to consist solely of
parthenogenic species. In Cryptostigma, most species appear to reproduce asexually, although a few species reproduce
sexually since immature males were observed, i.e. C. mexicanum, and in the rhizophilous species C. rhizophilum and C.
silveirai.
Cryptostigma Ferris
Cryptostigma Ferris 1922: 160. Type species: Cryptostigma ingae Ferris, by original designation and monotypy.
[=Cryptostigma inquilina (Newstead)]
Generic description. Adult female. Insect body oval to elongate oval, convex, rarely cylindrical when found
inside narrow hollow twigs, with a thin glassy wax cover or rarely a rather thicker wax cover. Not producing
an ovisac.
Dorsum. Dorsal derm generally becoming heavily sclerotized at maturity. Dorsal setae present or absent,
when present, spinose with a pointed, blunt, knobbed, rounded, or occasionally spatulate apex. Dorsal
tubercles absent. Sclerotic pores present or absent. Preopercular pores present or absent. Dorsal microducts
present, with either a single or bilocular opening, outer and inner ductules usually short. Simple pores usually
present, of 1 or more sizes, absent in Cryptostigma reticulolaminae. Orbicular pores present or absent; when
present, each pore with a thin membrane and margins membranous or mildly to heavily sclerotized;
sometimes with associated setae, simple pores and microducts. Cribriform plates absent, although cribriform
platelets often present. Anal plates together quadrate, rarely pyriform, with rounded angles, 4–21 setae on
dorsal surface, and 0–6 ventral subapical setae. Anal ring commonly with 10 setae, but with up to 20 setae in
C. serratum. Eyes absent.
Margin. Marginal setae spinose or conical, stout, with pointed apex, usually numerous, but absent in some
species. Stigmatic clefts deep; spiracular sclerotizations closely associated with a spiracle; spiracular
sclerotizations either short or long and extending far from margin, often enclosing spiracles. Stigmatic setae
totalling 0–3 per stigmatic area, present on each stigmatic sclerotization, setae often broken-off; shape bluntly
or sharply spinose, or conical, all subequal in length.
Venter. Spiracles large; width of peritreme usually greater than length of legs; spiracular opening
generally facing dorsally or towards body margin. Antennae reduced, usually 1–4 segmented, mostly 1
segmented or represented by a flattened segment bearing numerous setae, rarely up to 8 fused segments in C.
melissophilum. Legs greatly reduced; segments usually indistinct or fused; in many species represented by
clusters of setae usually associated with a tiny sclerotic plate. Spiracular disc-pores with 3–9 loculi.
Mouthparts well developed; labium with 8 labial setae. Pregenital disc-pores with 3–11 loculi, mostly with 5–
8 loculi, about same size or larger than spiracular disc-pores. Distribution of pregenital disc-pores variable.
Ventral setae, slender, pointed, usually abundant towards posterior end of body. Ventral tubular ducts usually
absent, but present in C. saundersi and C. chacoensis.
First-instar nymph. Body elongate oval.
Dorsum. Dorsal derm membranous, smooth or reticulated, with segmentation delineated by membranous
folds. Dorsal setae present or absent. Trilocular pore present on each side of head near margin close to each
eye. Dorsal microducts present. Simple pores present. Anal plates together oval or quadrate, dorsal surface
with 1 seta on anterior part of plate, often present on inner margin of plate, plus 3 apical setae; ventral side
with 1 fringe seta. Anal ring typical of coccid first-instar nymphs, with 6 setae and an irregular row of
translucent wax pores. Eyespots present or absent.
Margin. Outline smooth. Marginal setae sharply spinose, straight or strongly bent. Stigmatic setae bluntly
spinose, usually 1 or 2 per stigmatic cleft, but rarely up to 5.
Zootaxa 2709 © 2010 Magnolia Press · 9
REVISION OF CRYPTOSTIGMA FERRIS
Venter. Ventral derm membranous. Antennae 5 or 6 segmented, third antennal segment longest; if 5
segmented, fleshy setae present on last 2 segments only; if 6 segmented, fleshy setae present on last 3
segments. Interantennal setae usually 1 pair, rarely 3 pairs. Mid-ventral setae slender, usually present on last 3
abdominal segments, but may be present on all segments and mid-thoracic region. Submarginal setae present
in 7 pairs on each side between posterior stigmatic areas and anal cleft; with 1 or numerous setae between
anterior and posterior spiracles; with usually 1, rarely 2 pairs near apex of head; setae rarely present
elsewhere. Ventral microducts present submarginally from just below eyes to body posterior apex, rarely
present between pro- and metathoracic coxae and around mouthparts. Spiracular disc-pores with 3–9 loculi,
with 5-locular pores most common. Muscle plate usually attached to upper edge of each peritreme or absent.
Mouthparts well developed, with long coiled stylets (not illustrated in the figures); labium with 8 setae. Legs
well developed, often with numerous setae; microctenidia present on apex of tibiae. Prothoracic tarsal
digitules dissimilar, 1 knobbed and the other spiniform; meso- and metathoracic tarsal digitules similar,
knobbed. Claw slender, with or without a denticle; claw digitules slender, knobbed, one digitule broader than
other.
Diagnosis. Adult female. Without subcircular groups of simple disc pores present submarginally or
submedially, although a subcircular group may occur laterad to each spiracular sclerotization. Anal plates
together quadrate or subcircular. Pregenital disc pores present, either restricted to around vulva or extending
onto anterior abdominal segments; if pregenital disc-pores restricted to small area around vulva, then
stigmatic setae either absent or less than 3; ventral microducts short, without swollen inner ductules (except
for C. reticulolaminae Morrison); legs reduced, often represented by a group of setae and a small sclerotic
plate, never completely absent, although often hard to detect.
First-instar nymphs. Antennae 5 or 6 segmented; without a seta present near each coxa; with 1 or 2
membranous folds just above anal plates and always showing signs of sclerotization; usually with 3 ventral
submedian setae, rarely up to 5, or many setae present on all abdominal segments and thorax, never in 6 pairs
(known from the Neotropical region).
Key to the adult females of Cryptostigma Ferris
1. Ventral tubular ducts present.........................................................................................................................................2
- Ventral tubular ducts absent .........................................................................................................................................3
2. With multilocular disc-pores present just posterior to antennae; with 60 or more orbicular pores; associated with ants
............................................................................................................................................................. saundersi Laing
- Without multilocular disc-pores around antennae; with less than 35 orbicular pores; associated with stingless bees ..
............................................................................................................................................chacoensis Kondo, sp. nov.
3. Margin of dorsal stigmatic sclerotization dentate..................................................................serratum Kondo, sp. nov.
- Margin of dorsal stigmatic sclerotization smooth, not dentate.....................................................................................4
4. Dorsum with orbicular pores ........................................................................................................................................5
- Dorsum without orbicular pores.................................................................................................................................10
5. Dorsum with 2 orbicular pores; with multilocular disc-pores present on each side of mouthparts ...............................
......................................................................................................................................................biorbiculus Morrison
- Dorsum with 3 or more orbicular pores; without multilocular disc-pores around mouthparts ...................................6
6. Dorsum with a small group of numerous setae present next to each stigmatic sclerotization; dorsal derm becoming
tessellated in matured specimens..........................................................................................philwardi Kondo, sp. nov.
- Dorsal setae entirely absent; dorsal derm not tessellated in mature specimens............................................................7
7. Dorsum with 5 orbicular pores ........................................................................................................ urichi (Cockerell)
- Dorsum with 3 orbicular pores.....................................................................................................................................8
8. Body margin heavily papillated, with very few or no multilocular pores near margin of abdominal apex...................
............................................................................................................................................... gullanae Kondo, sp. nov.
- Body margin not papillated, with numerous multilocular pores present around submargin of abdomen....................9
9. Body about 2 times longer than wide; with 1 stigmatic seta per cleft; associated with ants..........................................
............................................................................................................................................ jonmartini Kondo, sp. nov.
KONDO
10 · Zootaxa 2709 © 2010 Magnolia Press
- Body about as long as wide; with 3 stigmatic setae per cleft; associated with stingless bees........................................
...................................................................................................................................... melissophilum Kondo, sp. nov.
10. Dorsum with sclerotic pores.......................................................................................................................................11
- Dorsum without sclerotic pores..................................................................................................................................16
11. Sclerotic pores arranged in square- to rectangular-shaped groups; outer ductule of dorsal microducts long and heav-
ily sclerotized, swollen near duct opening.............................................................................reticulolaminae Morrison
- Sclerotic pores not arranged as above; outer ductule of dorsal microducts long or short, if long, not swollen near duct
opening nor heavily sclerotized..................................................................................................................................12
12. Stigmatic sclerotization cone-shaped; dorsal surface of each anal plate with 10–16 setae....guadua Kondo & Gullan
- Stigmatic sclerotization not cone-shaped; dorsal surface of each anal plate generally with 5 or fewer setae............13
13. Tubercle-like preopercular pores present on mid dorsum and around anal plates ..........tuberculosum Kondo, sp. nov.
- Preopercular pores usually absent, if present, not tubercle-like.................................................................................14
14. Dorsum with small and subcircular groups of large simple pores next to stigmatic clefts ............................................
......................................................................................................................................... rhizophilum Kondo, sp. nov.
- Dorsum without small groups of simple pores next to stigmatic clefts......................................................................15
15. Venter with simple pores present on thorax............................................................................longinoi Kondo, sp. nov.
- Venter without simple pores ............................................................................................................. silveirai (Hempel)
16. Dorsum with clusters of 4–10 small simple pores; apex of some antennal fleshy setae always knobbed.....................
................................................................................................................................................... inquilinum (Newstead)
- Dorsum without clusters of simple pores; apex of antennal fleshy setae not knobbed, although rarely 1 or 2
branched.............................................................................................................................mexicanum Kondo, sp. nov.
Key to the known first-instar nymphs of Cryptostigma Ferris
1. Antennae 5 segmented..................................................................................................................................................2
- Antennae 6 segmented..................................................................................................................................................7
2. Dorsal setae in 2 submedian longitudinal rows............................................................................................................3
- Dorsal setae in 4 longitudinal rows.................................................................................tuberculosum Kondo, sp. nov.
3. Mid-ventral setae present on all abdominal and thoracic segments; marginal setae in a double row of, those on dorsal
surface sharply spinose, those on ventral surface flagellate ...................................................guadua Kondo & Gullan
- Mid-ventral setae present on last 3 abdominal segments only; marginal setae in a single row, all sharply spinose....4
4. With 2 pairs of ventral submarginal setae on head region between eyes; trochanter without a very long setae ...........
......................................................................................................................................................C. silveirai (Hempel)
- With 1 pair of ventral submarginal setae on head region between eyes; trochanter with a very long setae.................5
5. Femur of each leg with 6 to 8 setae (rarely 4 or 5).................................................................longinoi Kondo, sp. nov.
- Femur of each leg with 3 setae .....................................................................................................................................6
6. Ventral submarginal setae on abdomen of 2 types: outer submarginal setae sharply spinose, inner submarginal setae
slender; most marginal setae strongly bent...................................................................... rhizophilum Kondo, sp. nov.
- Ventral submarginal setae on abdomen all slender; marginal setae straight, not strongly bent......................................
.............................................................................................................................................. reticulolaminae Morrison
7. Dorsal setae absent........................................................................................................................biorbiculus Morrison
- Dorsal setae present......................................................................................................................................................8
8. Antennal segment III 2 or more times longer than antennal segment IV; femur with 4 to 6 setae ..............................9
- Antennal segment III only slightly longer than antennal segment IV; femur with 9 or more setae...............................
............................................................................................................................................chacoensis Kondo, sp. nov.
9. With 1 pair of ventral submedian setae between meso- and metathoracic area .........................................................10
- Without a pair of ventral submedian setae between meso- and metathoracic area ....................................................11
10. Femur with 5 setae; with no more than 1 microduct between each pair of submarginal setae on abdomen..................
............................................................................................................................................ jonmartini Kondo, sp. nov.
- Femur with 4 setae; with 3 to 4 microducts between most pairs of submarginal setae on abdomen.............................
...................................................................................................................................... melissophilum Kondo, sp. nov.
11. Claw denticle present; femur with 4 setae; anal plates smooth........................................................ .urichi (Cockerell)
- Claw denticle absent; femur with 4 or 6 setae; anal plates reticulated...................................... inquilinum (Newstead)
Zootaxa 2709 © 2010 Magnolia Press · 11
REVISION OF CRYPTOSTIGMA FERRIS
Description of species
Cryptostigma biorbiculus Morrison
(Figs 1E, 2, 3).
Cryptostigma biorbiculus Morrison, 1929: 48.
Cryptostigma biorbiculatus Morrison [misspelling], Williams 2001 (1999): 226.
Type material examined. Holotype. Adult . PANAMA (AS CANAL ZONE): Ancon, 5.v.1923, coll. W.M.
Wheeler (#110), ex Cordia alliodora, holotype clearly indicated on slide 1(2) (USNM). Paratypes. PANAMA
(AS CANAL ZONE): Corozal, 12.vii.1924, coll. W.M. Wheeler, ex Cordia alliodora, AL-098-84, 4 (4: 3
adult
+ 1 first-instar nymph) (AUCC); Corozal, 12.vii.1924, coll. W.M. Wheeler, ex Cordia alliodora, AL-
140-77, 2(2) (AUCC); Corozal, 12.vii.1924, coll. W.M. Wheeler, ex Cordia sp., AL-190-1999, slide mounted
from USNM dry material no. 666, 3(3) (AUCC).
Other material examined. COLOMBIA: Valle, Cali, 5.i.2000, coll. T. Kondo, ex Croton gossypiifolius,
tended by Azteca sp., AL-099-2000, 8 (8) (AUCC); Valle, Cali, 5.i.2000, coll. T. Kondo, ex Croton
gossypiifolius, tended by Azteca sp., AL-099-2000, 15 (15 first-instar nymphs) (AUCC); COSTA RICA:
Puntarenas, Parque Nacional Piedras Blancas (8°42’N, 83°12’W), 15.ii.2001, coll. M. Schmidt, ex
Tetrathylacium macrophyllum, tended by Azteca nigricans, 7 (7: 4 adult
+ 1 second-instar nymph + 2 first-
instar nymphs) (BME); PANAMA: Chiriqui Province, 1938, coll. Ferris, C. biorbiculus Morr., det T. Kondo,
no. 118, 1 (1 first-instar nymph) (BME).
Adult female (Figs 1E, 2)
Unmounted material. Alcohol-preserved specimens creamy yellow to orange, old specimens becoming
dark purplish red. Insects covered by a thin layer of flaky wax. Orbicular pores clearly visible as darker spots;
area around anal plates and stigmatic clefts darker; with a white wet wax around spiracular openings of
stigmatic cleft.
Mounted material. Body outline oval to elongate oval, 4–8 mm long, 3–6 mm wide (n=19).
Dorsum. Derm membranous, with numerous sclerotic pores of various geometric shapes (see Remarks),
scattered evenly on dorsum; each pore 8–31
m wide, with a central bilocular microduct, duct-rim of each
microduct about 4
m wide. Dorsal setae and preopercular pores absent. Simple pores each 3.5–6.0 m wide,
scattered evenly throughout dorsum. With 2 large subcircular orbicular pores, each 190–320
m wide at
widest point, 1 located dorsad to mouthparts, other on mid-thoracic region. Cribriform platelets each 4–17
m
wide, present singly or in groups of up to 40 platelets; with a group located on anterior margin of sclerotic area
around anal plates, and 4–6 groups present submedially between anal plates and mid-dorsal orbicular pore.
Anal plates together quadrate, each plate 156–189 mm long, 70–86
m wide; anterolateral margin 125–145
µm long, posterolateral margin 125–135 µm long; with about 10 setae on dorsal surface; plates located about
1/5 of body length from posterior margin. Anal ring with 8–10 setae. Eyes absent.
Margin. Body margin membranous, smooth, rugose or often papillated. Marginal setae each 13–20
m
long, sharply spinose, arranged in a single row, with 25–50 setae between each anterior and posterior
stigmatic areas. Stigmatic clefts deep, stigmatic sclerotization well developed, covering spiracles. Stigmatic
setae each 7.5–17.5 µm long and bluntly spinose, with 3–9 in each cleft but often broken off.
Vent er. Ventral setae slender, straight or slightly bent, each 17–32
m long, those on abdominal segments
longer. Ventral microducts each about 2
m wide, scattered evenly on venter. Tubular ducts absent.
Clypeolabral shield 253–286
m wide. Pregenital disc-pores each 5.0–6.0 m wide with 3–8 (mostly 7 & 8)
loculi; present on all abdominal segments and in mid-ventral areas of meso- and metathorax, with a denser
line of pores spreading from vulva to posterior spiracles. Spiracular disc-pores each 5.0–6.0
m wide, with 5–
8 (mostly 7 & 8) loculi. Antennae reduced, 1- or 2-segmented, total length 30–68
m long. With 3 or 4 pairs
of interantennal setae. Legs greatly reduced, with all segments fused, total length 53–85
m; tarsal digitules
similar, knobbed, but knobbed tips often broken off; claw digitules similar, knobbed. Claw denticle present or
absent. Each anterior spiracular peritreme 102–162
m wide, each posterior peritreme 119–162 m wide.
KONDO
12 · Zootaxa 2709 © 2010 Magnolia Press
FIGURE 1. Insects in life. A, colony of C. melissophilum inside nest of Schwarzula coccidophila (Apidae: Meliponini);
B, colony of C. chacoensis inside nest of stingless bees (Apidae: Meliponini); C, C. chacoensis; D, C. reticulolaminae;
E, C. biorbiculus; F, C. guadua, G, C. longinoi, H, C. inquilinum; I, C. serratum; J, C. jonmartini; K, C. urichi.
Zootaxa 2709 © 2010 Magnolia Press · 13
REVISION OF CRYPTOSTIGMA FERRIS
FIGURE 2. Cryptostigma biorbiculus Morrison, adult female.
KONDO
14 · Zootaxa 2709 © 2010 Magnolia Press
FIGURE 3. Cryptostigma biorbiculus Morrison, first-instar nymph.
Diagnosis. The adult female of C. biorbiculus can be diagnosed by the following features: (i) dorsum with
numerous sclerotic pores, (ii) a pair of orbicular pores present in midline, (iii) 3 or more pairs of small groups
of cribriform platelets present submedially between anal plates and posterior orbicular pore, (iv) preopercular
Zootaxa 2709 © 2010 Magnolia Press · 15
REVISION OF CRYPTOSTIGMA FERRIS
pores absent, (v) 9–15 setae present on dorsal surface of each anal plate, (vi) ventral tubular ducts absent, and
(vii) multilocular disc-pores present near mouthparts.
Remarks. Specimens collected in Cali, Colombia, on Croton gossypiifolius have sclerotic pores with
smooth round edges, in contrast to more irregular to quadrate sclerotic pores as seen in the type specimens and
species collected elsewhere.
First-instar nymph (Fig. 3)
Unmounted material. Not available for study.
Mounted material. Elongate oval, 0.7–1.6 mm long, 0.4–0.7 mm wide (n=16).
Dorsum. Dorsal derm membranous, often reticulated, with segmentation delineated by membranous folds.
Dorsal setae absent. Trilocular pore present on each side of head near margin close to eye. Dorsal microducts
each about 2
m wide, numerous over dorsum. Simple pores each 3–4 m wide, with center of pore either
with a darker region or of normal type. Anal plates together oval, each plate 58–72
m long, 21–26 m wide;
dorsal surface with 1 seta on anterior part of plate plus 3 apical setae; ventral surface with 1 fringe seta. Anal
ring as in generic diagnosis. Eyespots present just above level of antennal scape.
Margin. Outline smooth. Marginal setae sharply spinose, total number 64–75, with 8–12 anteriorly
between eyes, and on each side 6–8 between each eye and anterior stigmatic setae, 6–8 between each group of
anterior and posterior stigmatic setae, and 14–16 between posterior stigmatic setae and anal cleft. Stigmatic
setae bluntly spinose, each seta 18–24
m long; with 1 or 2 (mostly 1) stigmatic setae in each anterior cleft
and 1–3 (mostly 2) in each posterior cleft.
Ven te r. Microspines present on all abdominal segments. Antennae 6 segmented, each 199–318
m long.
Mid-ventral setae slender, 1 pair present on last 3 abdominal segments. Interantennal setae 1 pair.
Submarginal setae in 7 pairs on each side between posterior stigmatic areas and anal cleft, 1 seta between each
anterior and posterior spiracles, and 1 pair near apex of head. Ventral microducts each about 2
m wide,
present submarginally from just posterior to eyes to posterior body apex. Spiracular disc-pores each 3.5–4.0
m wide, with 3–9 (mostly 5 & 7) loculi; each anterior spiracular furrow with 8–11 pores; each posterior
spiracular furrow with 11–14 pores. Spiracular peritremes each 15–16
m wide. Clypeolabral shield 79–164
m wide. Legs well developed, with numerous setae; trochanter + femur 270–361 m long, tibia + tarsus
345–426
m long. Prothoracic tarsal digitules dissimilar, 1 knobbed and other spiniform; meso- and
metathoracic tarsal digitules similar, knobbed. Claw slender, with or without a denticle.
Diagnosis. The first-instar nymph of C. biorbiculus can be diagnosed by the following features: (i) dorsal
setae absent, (ii) each anterior stigmatic cleft with 1 or 2 stigmatic setae; each posterior cleft with 1–3 setae,
(iii) 6–8 marginal setae present between anterior and posterior stigmatic setae, (iv) antennae 6 segmented, (v)
with 3 pairs of ventral submedian setae on abdomen, and (vi) each anterior spiracular furrow with 8–11 pores
and each posterior furrow with 11–14 pores.
Remarks. First-instar nymphs from material collected in Colombia on Croton gossypiifolius have many
fewer setae on the legs, a greater number of ventral and dorsal microducts and a small apical claw denticle
(denticle absent in type material).
Host plants. Boraginaceae: Cordia alliodora, Cordia sp.; Euphorbiaceae: Croton gossypiifolius;
Flacourtiaceae: Tetrathylacium macrophyllum.
Associated Hymenoptera. Dolichoderinae: Azteca longiceps, A. nigricans, Azteca sp.;
Pseudomyrmecinae: Pseudomyrmex sericea (as Pseudomyrma sericeae) (Morrison, 1929).
Males. Unknown.
Distribution. Neotropical Region: Colombia, Costa Rica, Panama.
Cryptostigma chacoensis Kondo, sp. nov.
(Figs 1B, C, 4 & 5)
Type material examined. Holotype. Adult
. BOLIVIA: Santa Cruz, Charagua, 50 km N.E. La Brecha, -
.xi.2002, coll. D.W. Roubik, ex Caesalpinia paraguariensis, 1 (1) (USNM). Paratypes. Same data as holotype,
KONDO
16 · Zootaxa 2709 © 2010 Magnolia Press
7 (13: 4 adult +1 third-instar nymph + 8 embryonic nymphs) (AUCC); 6 (9: 4 adult + 1 third-instar
nymph + 4 embryonic nymphs) (BME); 6 (9: 4 adult
+ 1 third-instar nymph + 4 embryonic nymphs)
(BMNH); 21 (38: 4 adult
+ 1 adult anal plate + 5 third-instar nymphs + 2 second-instar nymphs + 26
first-instar nymphs including embryonic nymphs) (USNM).
Adult female (Figs 1B, C & 4)
Unmounted material. Insects found inside nests of Plebeia sp. and Schwarzula timida Camargo and
Pedro (D.W. Roubik, pers. comm.). Young adult females reddish orange, becoming purplish brown at
maturity; anal plates and surrounding area distinctly darker than rest of dorsum.
Mounted material. Body outline oval to elongate oval, constricted at stigmatic areas, 4.8–6.9 mm long,
4.5–6.1 mm wide (n=17).
Dorsum. Derm membranous, with 25–33 subcircular, compound orbicular pores present submarginally
and submedially, absent from mid-dorsum; diameter of each pore 50–350 (mostly 100–200)
m wide, those
closest to body margin usually smaller; present submarginally and submedially, absent from mid-dorsum;
each orbicular pore often associated with very small simple pores, and often with a few bilocular microducts
arranged near the sclerotized orbicular pore rim; each pore with 30–50 slender setae, each 15–28
m long,
around pore margins, either on sclerotic rim or on membrane near rim, rarely present on central area of pore.
Dorsal setae absent, except for those on orbicular pores. Simple pores with a pore rim of variable thickness,
each 5–7
m wide. Dorsal microducts with a bilocular opening, of 2 types, duct-rim of small simple ducts
each about 2
m wide, duct-rim of larger ducts each 5 m wide. Preopercular pores absent. Narrow sclerotic
crescent present around anal plates. Anal plates heavily sclerotized, plates together quadrate, with rounded
angles, each plate 235–260
m long, 90–125 m wide, anterolateral margin 135–180 m long, posterolateral
margin 155–180
m long; setae not visible on most specimens, but with 18 setae on dorsal surface of 1
specimen; plates located about 1/5–1/4 of body length from posterior margin. Anal ring with 10 setae.
Eyespots absent.
Margin. Margins smooth. Marginal setae conical, basal portion of seta often swollen, with well-developed
setal sockets, each seta 10–33
m long, numerous, with 30–90 setae in 1 or 2 rows between each anterior and
posterior stigmatic areas. Stigmatic clefts deep, forming a sclerotic plate enclosing spiracles, each cleft with 3
bluntly spinose stigmatic setae, straight or curved, subequal in length, each 20–43
m long, but often broken
off.
Venter. Derm membranous. Ventral setae slender, those on abdominal segments longer (each 20–65
m
long), shorter around body margin (13–35
m long). Ventral microducts each 2–4 m wide, each showing
slight sclerotization around outer rim; evenly distributed on venter. Tubular ducts present around vulva and
anteriorly on mid-areas of abdomen up to about abdominal segment IV. Clypeolabral shield 480–560
m
wide. Multilocular disc-pores each 5–9
m wide, with 6–8 (mostly 7 or 8, rarely with 4, 5 or 9) loculi, present
around vulva and medially on all abdominal segments; also present transversely on thorax, with pores
extending to mesothoracic legs and posterior spiracles. Spiracular disc-pores each 5–7
m wide with 5–7
(mostly 6, rarely with 4 or 5) loculi. Antennae reduced, 1–4 segmented, total length 65–190
m. Legs
reduced, femora, tibiae and tarsi strongly fused, total length 65–200
m long; tarsal digitules setose; claw
digitules knobbed. Claw denticle absent. Spiracles located between margin and each prothoracic and
mesothoracic legs; each anterior peritreme 190–215
m wide, each posterior peritreme 190–240 m wide.
Diagnosis. The adult female of C. chacoensis can be diagnosed by the following combination of
characters: (i) dorsum with 25–33 complex orbicular pores, (ii) unilocular and bilocular dorsal microducts
present, (iii) with about 18 setae on surface of each anal plate, (iv) preopercular pores absent, (v) marginal
setae sharply spinose, often with swollen bases, and (vi) presence of ventral tubular ducts. C. chacoensis
appears most similar to C. saundersi Laing, but the following features separate the two species (characters on
C. saundersi in brackets): (1) only 22–23 complex orbicular pores present (60 or more); (2) multilocular pores
absent near antennae (a multilocular pore present next to each antenna); and (3) ventral microducts arranged
randomly (arranged in transverse rows).
Zootaxa 2709 © 2010 Magnolia Press · 17
REVISION OF CRYPTOSTIGMA FERRIS
FIGURE 4. Cryptostigma chacoensis Kondo, adult female.
KONDO
18 · Zootaxa 2709 © 2010 Magnolia Press
FIGURE 5. Cryptostigma chacoensis Kondo, first-instar nymph.
Remarks. Cryptostigma chacoensis together with C. melissophilum are the only species of Coccidae
known to be tended by stingless bees and harboured within their nests. Camargo and Pedro (2002a) reported
Zootaxa 2709 © 2010 Magnolia Press · 19
REVISION OF CRYPTOSTIGMA FERRIS
in detail on the association between the stingless Schwarzula sp. bees and C. melissophilum (then
Cryptostigma sp.), and later described the bee as Schwarzula coccidophila (Camargo & Pedro, 2002b).
First-instar nymph (Fig. 5)
Unmounted material. Immature insects yellowish.
Mounted material. Elongate oval, 1.0–1.1 mm long, 0.5–0.6 mm wide (n=41).
Dorsum. Dorsal derm membranous, highly rugose, with segmentation delineated by membranous folds.
Dorsal setae present in 2 mid-dorsal longitudinal rows, each seta 7–10
m long. Trilocular pore present on
each side of head near margin close to eye. Dorsal microducts not detected. Simple pores about 3
m wide,
present in about 3 longitudinal rows, most pores in pairs. Anal plates triangular, each plate 80–83
m long,
23–33
m wide; dorsal surface with 1 seta on margin of mid part of plate plus 3 apical setae; ventral surface
with 1 fringe seta. Anal ring with 6 setae and an irregular row of translucent wax pores. Eyespots present just
above level of antennal scape.
Margin. Outline smooth. Marginal setae sharply spinose, each 12–18
m long; total number 76–93: with
10–15 anteriorly between eyes, and on each side, 7–10 between each eye and anterior stigmatic setae, 8–10
between each anterior and posterior stigmatic setae, and 16–20 between posterior stigmatic setae and anal
cleft. Stigmatic setae bluntly spinose, each 17–33
m long, with 1 or 2 per stigmatic cleft.
Venter. Ventral derm membranous; microspines present on mid-ventral and mid-thoracic areas (not
illustrated). Antennae 6 segmented, total length 285–300
m. Mid-ventral setae slender, with 1 pair present on
last 5 abdominal segments. Interantennal setae 1 pair. Submarginal setae present in 7 pairs on each side of
abdomen (a pair in each segment), a single seta between each anterior and posterior spiracles, and 1 pair near
apex of head. Ventral microducts each about 2
m wide, 1 present mesad to each antennal scape, 1 on mid-
area between antennal scape and anterior stigmatic setae, 1 between each anterior and posterior stigmatic
setae, and 6 next to first 6 pairs of submarginal setae on abdomen on each side, just mesad to each inner
submarginal seta. Spiracular disc-pores each 4–5
m wide with 3–8 (mostly 5 & 7) loculi; each anterior and
posterior spiracular furrow with 6–9 pores. Spiracular peritremes each 12–15
m wide. Clypeolabral shield
100–110
m wide. Legs well developed, with numerous setae; trochanter + femur each 170–180 m long,
tibia + tarsus each 250–260
m long, microctenidia not visible on tibial apex. Tarsal digitules similar,
knobbed. Claw slender, without a denticle; claw digitules slender, knobbed, 1 slightly thicker than other.
Diagnosis. The first-instar nymph of C. chacoensis can be diagnosed by the following features: (i) dorsal
derm rugose, (ii) dorsal setae present in 2 longitudinal submedian rows of about 11 pairs of setae, (iii)
stigmatic setae totalling 1 or 2 per stigmatic cleft, (iv) 8–10 marginal setae present between anterior and
posterior stigmatic setae, (v) antennae 6 segmented, (vi) with 5 pairs of ventral submedian setae on abdomen,
(vii) spiracular disc-pores with 3–8 loculi, and (vii) each anterior and posterior spiracular furrow with 6–9
pores.
Host plants. Fabaceae: Caesalpinia paraguariensis.
Associated Hymenoptera. Meliponini: Schwarzula timida and Plebeia sp. (D. Roubik, pers. comm.).
Males. Unknown.
Etymology. The species is named after the biodiversity hotspot, the Chaco region of Bolivia, where it was
collected.
Distribution. Neotropical Region: Bolivia.
Cryptostigma guadua Kondo & Gullan
(Figs 1F, 6 & 7)
Cryptostigma guadua Kondo & Gullan, 2004: 717.
Type material examined. Holotype. Adult . PERU: Madre de Dios, Parque Nacional Manu, Estación
Biológica de Cocha Cashu, 11°54’S, 71°22’W, c. 350 m. a.s.l., -.vi.2003, coll. D.W. Davidson, ex bamboo
(Guadua sp.), inside nest of Camponotus mirabilis, 1 (1) (MUSM). Paratypes. Same data as holotype 13 (13:
3 adult
+ 5 second-instar nymphs + 5 first-instar nymphs) (BME, UNAD, USNM); same data as holotype
but c. 400 m. a.s.l., 8.x.2001, No. cc-01-120, 3 (3: 3 first-instar nymphs) (UNAD, USNM); same data as
holotype but 31.v.2003, No. 03-043, tended by Pseudocolobopsis sp. 3 (3: 2 adult
+ 1 third-instar nymph)
(BME, UNAD); same data as holotype but 11°54’S, 71°22’W, c. 400 m. a.s.l., 13.xi.2001, No. cc-01-140 &
KONDO
20 · Zootaxa 2709 © 2010 Magnolia Press
cc-01-157, inside nest of Camponotus longipilis, 4 (4) (UNAD); Tambopata Research Center on Rio
Tambopata, -.ii.2002, coll. D.W. Davidson, No. 01-TRC-01, ex bamboo, 2 (2: 1 adult female + 1 pharate adult
) (UNAD).
FIGURE 6. Cryptostigma guadua Kondo & Gullan, adult female.
Zootaxa 2709 © 2010 Magnolia Press · 21
REVISION OF CRYPTOSTIGMA FERRIS
Adult female (Figs 1F & 6)
Unmounted material. Color variable, live specimens often reddish with a yellow tinge. Dorsum of
specimens preserved in alcohol pale orange to yellowish-brown, ventral surface pinkish. Dorsal surface
covered by a thin clear waxy layer, easily detached in alcohol, wax of a flaky texture. Stigmatic cleft and anal
cleft very deep; stigmatic sclerotizations and anal plates clearly marked and light brown to dark brown in
color.
Mounted material. Body outline broadly oval, margins smooth or irregular, 3.8–10.1 mm long, 4.0–9.0
mm wide (n=12).
Dorsum. Derm membranous, with numerous circular or constricted sclerotic pores; abundant towards
body margin; each pore 13–100
m wide, sclerotic pores near body margin generally larger and often with
constricted margins; density of sclerotic pores highly variable between specimens, ranging from very scarce
to dense, but always fewer on mid dorsum. Dorsal setae each 7–18
m long, sharply or bluntly spinose, each
seta on a small dorsal sclerotic plate, each 5–18
m wide; present evenly on dorsum. Preopercular pores each
about 10–15
m wide, tubercle-like, present on mid-dorsum anterior and laterad to anal plates. Dorsal
microducts each about 3
m wide, numerous, scattered on dorsum, with a very long and usually slightly
sclerotized outer ductule; those near margin often longer; inner ductule about as long as or longer than outer
ductule. Sclerotic crescent present around anal plates. Anal plates together pyriform, occasionally quadrate,
with rounded angles; each plate 133–183
m long, 55–80 m wide, anterolateral margin 103–145 m long,
posterolateral margin 75–103
m long; with 10–16 setae on dorsal surface of each plate; plates located about
1/3–1/4 of body length from posterior margin. Anal ring with 10 setae. Eyes absent.
Margin. Marginal setae not differentiated. Stigmatic clefts very deep, forming a conical stigmatic
sclerotization; each cleft 480–680
m long, opening 270–460 m wide. Stigmatic setae usually hard to detect,
although 3 short conical swellings often present on outer margin of stigmatic sclerotization suggesting the
presence of stigmatic setae.
Ven te r. Derm membranous. Ventral setae slender, straight or slightly bent, those on abdominal segments
usually longer, each 20–43
m; those near margins shorter, each 7–15 m long. Interantennal setae about 3
pairs. Ventral microducts each about 3
m wide, scattered evenly on venter. Tubular ducts absent.
Clypeolabral shield 305–335
m wide. Multilocular disc-pores each 7–10 m wide with 3–8 (mostly 6 or 7)
loculi, abundant around vulva and present on all abdominal segments; pores also present around metathoracic
legs in a line to posterior spiracle. Spiracular disc-pores each 6–8
m wide with 3–8 (mostly 5) loculi, some
extending anteriorly towards mesothoracic legs. Antennae very small, 45–65
m long, apparently 1
segmented, represented by a flattened segment bearing about 14–15 setae. Legs greatly reduced, represented
by a rudimentary claw, several setae, and associated pores, total length 45–65
m. Spiracles much larger than
legs, located far from body margin; each anterior spiracular peritreme 113–143
m wide, each posterior
peritreme 125–160
m wide.
Diagnosis. The adult female of C. guadua can be diagnosed by the following features: (i) dorsum with
numerous small sclerotic pores, (ii) presence of a deep, cone-shaped stigmatic sclerotization, (iii) presence of
tubercle-like preopercular pores, (iv) with 10–16 setae on dorsal surface of each anal plate, (v) multilocular
disc-pores present on all abdominal segments, surrounding the spiracles and including a group which forms a
line from each mesothoracic leg to the posterior spiracle, and (vi) absence of ventral tubular ducts.
First-instar nymph (Fig. 7).
Unmounted material. Not available for study.
Mounted material. Elongate oval; recently eclosed or early settled nymphs 1.2–1.6 mm long, 0.9–1.4
mm wide (n=4), fully-grown nymphs 1.8–2.2 mm long, 1.6–1.9 mm wide (n=4).
KONDO
22 · Zootaxa 2709 © 2010 Magnolia Press
FIGURE 7. Cryptostigma guadua Kondo & Gullan, first-instar nymph.
Dorsum. Dorsal derm membranous, tesselated on some specimens, with segmentation delineated by
membranous folds. Dorsal setae present in 2 submedian parallel rows, each seta about 15
m long, setae on
Zootaxa 2709 © 2010 Magnolia Press · 23
REVISION OF CRYPTOSTIGMA FERRIS
head region usually present, other dorsal setae often broken off or undetectable. Trilocular pore present on
each side of head near margin close to eye. Dorsal microducts each about 2
m wide, numerous over dorsum.
Simple pores about 3–5
m wide, those closer to body margin slightly smaller. Anal plates together pyriform,
each plate 165–180
m long, 70–80 m wide, anterolateral margin 138–163 m long, posterolateral margin
48–85
m long; dorsal surface of shingled texture, with 1 seta on anterior part of plate plus 4 apical setae;
ventral surface with 1 fringe seta. Anal ring as in generic diagnosis.
Margin. Outline smooth. Marginal setae of 2 types; dorsal marginal setae sharply spinose, total number
133–160, with 15–20 anteriorly between eyes, and on each side, 10–15 between each eye and anterior
stigmatic setae, 16–23 between each group of anterior and posterior stigmatic setae, and 29–49 between
posterior stigmatic setae and anal cleft; ventral marginal setae flagellate, more numerous than dorsal marginal
setae, each 63–90
m long.
Ve nt er. Ventral derm membranous, with microtrichia present on abdominal segments. Mid-ventral setae
flagellate, each 75–100
m long, present on all segments posterior to mouthparts, with 4–8 setae per segment.
Submarginal ventral setae similar to ventral marginal setae, in 6 pairs on each side of abdomen, about 7 setae
between each anterior and posterior spiracle, and 1 seta between each antennal scape and anterior spiracular
furrow. Stigmatic setae each sharply spinose, short, each seta 15–20
m long, subequal in length; anterior
stigmatic setae in a group of 3 or 4 setae, posterior stigmatic setae in a group of 4 or 5 setae. Interantennal
setae in 3 or 4 pairs. Ventral microducts each 2–3
m wide, with a very long inner ductule; numerous around
body margin and in a transverse row posterior to mouthparts and hind coxae. Spiracular disc-pores each about
5
m wide with 5–7 (mostly 5)-loculi, with 12–18 pores per spiracular furrow. Clypeolabral shield 180–214
m wide. Legs well developed, with numerous setae; trochanter + femur 215–245 m long, tibia + tarsus
235–265
m long, microctenidia not visible on tibial apex. Tarsal digitules similar, spiniform. Claw slender,
without a denticle; claw digitules slender, with knobbed apices. Antennae 5 segmented, total length 390–440
m.
Diagnosis. The first-instar nymph of C. guadua can be diagnosed by the combination of the following
features: (i) dorsal setae arranged in 2 parallel longitudinal lines, (ii) numerous mid-ventral setae present on
all abdominal and thoracic segments, (iii) antennae 5 segmented, (iv) 2 types of marginal setae present, those
on dorsal surface sharply spinose, those on ventral surface flagellate, and (v) presence of numerous ventral
microducts arranged in a transverse row just posterior to the mouthparts and anterior coxae.
Remarks. The preceding description is adapted from Kondo and Gullan (2004), with copyright
permission from Neotropical Entomology.
Host plants. Poaceae: bamboo (Guadua sp.)
Associated Hymenoptera. Formicinae: Camponotus (Myrmostenus) longipilis, C. (M.) mirabilis and C.
(Pseudocolobopsis) sp.
Males. Unknown.
Distribution. Neotropical Region: Peru.
Cryptostigma gullanae Kondo, sp. nov.
(Fig. 8)
Material examined. Holotype. Adult
. ECUADOR: Provincia Napo, Jatun Sacha, 400 m asl, 1°04’S,
37°37’W, 3.viii.1991, coll. P.S. Ward (#11320), ex live branch of Sapium utile, tended by Pseudomyrmex
viduus, 1 (2: 1 adult
+ 1 third-instar nymph), holotype clearly indicated on label (USNM).
Adult female (Fig. 8)
Unmounted material. Not available for study.
Mounted material. Body outline irregularly oval, constricted at stigmatic areas, 3.1 mm long, 2.3 mm
wide (n=1).
KONDO
24 · Zootaxa 2709 © 2010 Magnolia Press
FIGURE 8. Cryptostigma gullanae Kondo, adult female.
Dorsum. Derm membranous, highly rugose; with 3 subcircular orbicular pores: 1 on head region, and 1 on
each side between anterior and posterior stigmatic areas; each pore 225–275
m wide. Dorsum with
Zootaxa 2709 © 2010 Magnolia Press · 25
REVISION OF CRYPTOSTIGMA FERRIS
conspicuous apodemes evenly distributed on dorsum; most conspicuous in submarginal areas. Dorsal setae
absent. Cribriform platelets each 5–10
m wide, with 3–6 pores, either square, hexagonal, triangular or
irregular in shape. Simple pores each 6–8
m wide, thick, sunken, scattered evenly on dorsum. Dorsal
microducts each about 3–4
m wide, with a simple opening, thick rimmed, sunken, evenly distributed on
dorsum. Preopercular pores absent. Sclerotic crescent present around anal plates. Anal plates together
quadrate, with rounded angles, each plate 205–220
m long, 75–88 m wide, anterolateral margin 148–150
m long, posterolateral margin 105–108 m long; with 10 or 11 setae on dorsal surface; plates located about
1/5 of body length from posterior margin. Anal ring with 6 setae. Eyes absent.
Margin. Body margin papillate. Marginal setae sharply spinose, each 15–20
m long, with well-developed
bases, arranged in a single irregular row, numerous, with about 50–60 between anterior and posterior
stigmatic areas. Stigmatic clefts very deep, stigmatic sclerotization well-developed, extending from margin
onto area dorsad to spiracles. Stigmatic setae similar to marginal setae, with 3 per cleft, subequal in length,
each 13–18
m long, often broken off.
Vent er. Derm membranous. Ventral setae slender, straight or slightly bent, each 15–35
m long, abundant
on abdominal segments and toward margin, longest anterior to vulva. Ventral microducts each 3–4
m wide,
scattered evenly on venter. Tubular ducts absent. Clypeolabral shield 203
m wide. Multilocular disc-pores
each 4–6
m wide with 6–8 (mostly with 5 and 6) loculi, sunken; present around vulva and on last abdominal
segments, becoming progressively fewer on anterior segments; pores absent medially and only present near
the areas of mediolateral lobes on segments V and IV. Spiracular disc-pores each 6–7
m wide, with 4–7
(mostly with 5 & 6) loculi, sunken, numerous, found laterad to each spiracular peritreme and incorporated into
stigmatic sclerotization. Antennae reduced, each with 1 or 2 antennal segments, fleshy setae normal, total
length 58–68
m. Legs reduced, atrophied, but claws well developed; with most segments fused, total length
38–50
m. Claw and tarsal digitules knobbed. Claw with or without a denticle. Spiracles larger than legs; each
anterior spiracular peritreme 175–180
m wide, each posterior peritreme 170–190 m wide.
Diagnosis. The adult female of C. gullanae can be diagnosed by the following combination of characters:
(i) dorsal setae absent, (ii) dorsum with 3 orbicular pores, (iii) cribriform platelets present, (iv) apodemes
conspicuous on dorsal derm, (v) dorsal surface of anal plates with 10 or 11 setae, (vi) preopercular pores
absent, (vii) margin papillate, (viii) with 3 stigmatic setae in each cleft, but often broken off, (ix) multilocular
disc-pores present around vulva and medially on last abdominal segments, but becoming progressively fewer
on anterior abdominal segments, and sparsely scattered submarginally on abdomen, and (x) anal ring with 6
setae.
Remarks. First-instar nymphs were not available during present study.
Host plants. Euphorbiaceae: Sapium utile.
Associated Hymenoptera. Formicidae: Pseudomyrmecinae: Pseudomyrmex viduus.
Males. Unknown.
Etymology. The species is name after Dr. Penelope J. Gullan who prepared the slide-mounts. Dr. Gullan is
an eminent scale insect taxonomist and expert in the study of ant-coccoid interactions.
Distribution. Neotropical Region: Ecuador.
Cryptostigma inquilinum (Newstead)
(Figs 1H, 9 & 10)
Pseudophilippia inquilina Newstead, 1920: 181. Lectotype , Jamaica (BMNH). [Lectotype by subsequent designation,
Qin & Gullan 1989: 225].
Akermes secretus Morrison, 1922: 145. Holotype
, Puerto Rico, on Inga laurina (USNM) [Synonymized by Qin &
Gullan, 1989: 225].
Cryptostigma ingae Ferris, 1922: 160. Holotype
, Puerto Rico, on Inga laurina (BME) [Synonymized by Qin &
Gullan: 1989: 225].
Cryptostigma inquilina (Newstead); Steinweden, 1929: 234. Change of combination.
Cryptostigma secretus (Morrison); Morrison, 1929: 53. Change of combination. [Synonymized by Qin & Gullan: 1989:
225]
KONDO
26 · Zootaxa 2709 © 2010 Magnolia Press
Type material examined. Holotype. Adult , Akermes secretus Morrison. PUERTO RICO: on Inga laurina
(USNM). Paratypes. PUERTO RICO: Lares, 21.i.1922, coll. not given, ex Inga laurina, 3 (4) (BME).
FIGURE 9. Cryptostigma inquilinum (Newstead), adult female.
Zootaxa 2709 © 2010 Magnolia Press · 27
REVISION OF CRYPTOSTIGMA FERRIS
Other material examined. BRITISH GUYANA: Camaria Falls, date not given, coll. I.W. Bailey, ex
Triplaris surinamensis, AL-271-75, labelled as C. secretus (Morr.), 3 (3) (AUCC); COSTA RICA: near Zent,
1924, coll. W.M. Mann, ex Inga laurina, labelled Akermes secretus, 1 (2) (USNM); CUBA: Cien Fuegos,
18.iv.1956, coll. P.A. Berry, ex algarrobo, #56-5866 (56-664) 1 (2) (USNM); GRENADA: 8.viii.1895, coll.
F.W.A., host not given, labelled C. urichi (Ckll.) secretus?, 2 (2) (USNM); MEXICO: Acapulco, Jardin
Botánico de Acapulco, Universidad Loyola del Pacífico, 16˚49'37.9"N, 99˚50'49.9"W, 15.i.2007, coll. T.
Kondo, ex Acacia paniculata, tended by Crematogaster sp., 16 (16) (BME); PUERTO RICO: Lares,
21.i.1922, coll. not given, ex Inga laurina, labelled C. ingae, 2 (2) (BME); 1921, coll. G.N. Wolcott, ex Inga
laurina, slide mounted from type material 6.i.1965, R.F. Wilkey, labelled Cryptostigma ingae Ferris 5 (8: 2
adult
+ 6 first-instar nymphs) (BME); Manati, 1923, coll. G.N. Wolcott, ex Ficus laevigata, labelled C.
inquilina (Newst.), 2 (2) (BME); Mayaguez, date not given, coll. W. Hoover 4 (8: 2 adult
+ 6 first-instar
nymphs) (USNM); Mayaguez, 30.iii.1915, coll. R.H. Yan Zwalowenburg, ex branches of Inga laurina, slide
mounted from type material, AL-075-86, labelled C. secretus Morrison, 8 (8) (AUCC); Mayaguez,
30.iii.1915, coll. R.H. Yan Zwalowenburg, ex branches of Inga laurina, slide mounted from type material,
AL-085-84, labelled C. secretus Morrison, 14 (39: 1 adult
+ 38 first-instar nymphs) (AUCC); Mayaguez,
let. 30.iii.1915, coll. R.H. Yan Zwalowenburg, ex branches of Inga laurina, AL-150-77, labelled C. secretus
Morrison, 6 (6) (AUCC); San Sebastian, 25.xi.1935, coll. M.R. Smith, ex Inga inga, AL-274-75, 4 (4)
(AUCC); Mayaguez, let. 30.iii.1915, coll. R.H. Yan Zwalowenburg, ex Inga laurina, labelled C. secretus
Morrison, 1 (4) (BME); Mayaguez, let. 30.iii.1915, coll. R.H. Yan Zwalowenburg, ex Inga laurina, labelled
A. secretus Morrison, 2 (11: 3 adult
+ 8 first-instar nymphs) (USNM); San Juan, 19.ix.1911, coll. W.V.
Tower, ex guama twigs (=Inga sp.), AL-270-75, labelled C. secretus Morrison, 4 (4) (AUCC); TRINIDAD &
TOBAGO: South Tobago, Friendship State, 7.xi.1918, coll. H. Morrison, ex Haematoxylon campechiamun,
slide mounted from USNM dry material A-912, AL-076-86, labelled C. secretus Morrison, 13 (13) (AUCC).
Adult female (Figs 1H & 9)
Unmounted material. “Color in life of different shades of light reddish or yellowish brown, often
varying to hint of lavender purple brown; color of alcoholic specimens dorsally pale grayish brown, with
somewhat distinct linear transverse mottling of dark brown, tiny blackish flecks along margin, brown area
around anal plates, small white spots at stigmatic area, ventrally with marginal band of dull brown, anterior 2
thirds of surface yellow cream, shading off to brown at edges, ventral abdominal segments about same color
as dorsum; whole venter variously mottled, flecked with dark color” (Morrison, 1922). Specimens collected in
Mexico by the author (Fig. 1H) with a rugose dorsum with clearly visible dorsal segmentation; color ranging
from yellow to pink, with anal plates darker, orange brown to red brown, with a conspicuous white wax on
each stigmatic area.
Mounted material. Body outline oval to irregularly oval, 1.8–7.0 mm long, 1.7–6.4 mm wide (n=60).
Dorsum. Derm membranous, becoming highly sclerotized at maturity. With 5 large to median-size,
subcircular orbicular pores: 1 on head region between antennae and apex of head, and 1 mesad to each
spiracular sclerotization. Dorsal setae flagellate, each 32–79
m long, with about 5–20 setae in a single row
around margins of each stigmatic sclerotization, completely absent from rest of dorsum. Simple pores of 2
types: large pores each about 5
m wide; numerous, scattered evenly on dorsum; small pores, each about 4 m
wide, present in clusters of 4–10 pores scattered on dorsum. Dorsal microducts bilocular, each about 3–4
m
wide, numerous, scattered evenly on dorsum. Preopercular pores absent. Sclerotic crescent present around
anal plates. Anal plates together oval, each plate 216–243
m long, 81–119 m wide, anterolateral margin
124–162
m wide, posterolateral margin 129–146 m wide; with 11–17 setae on dorsal surface of each plate,
plus about 3 ventral subapical setae on each plate; plates located about 1/5 of body length from posterior
margin. Anal ring with 10 setae. Eyes absent.
Margin. Margins smooth. Marginal setae sharply spinose, each 15–24
m long, with swollen bases,
arranged in a single row, numerous, with about 40–50 between anterior and posterior stigmatic areas.
Stigmatic clefts very deep, stigmatic sclerotization short, forming a sclerotized crescent incorporating many
multilocular pores. Stigmatic setae usually 2 (occasionally 3) per cleft, each 19–47
m long, bluntly spinose,
all subequal in length, but often broken off.
KONDO
28 · Zootaxa 2709 © 2010 Magnolia Press
FIGURE 10. Cryptostigma inquilinum (Newstead), first-instar nymph.
Vent er. Derm membranous. Ventral setae slender to sharply spinose, straight or slightly bent, those on mid
venter longest, each 34–53
m long; those close to margin shortest, 24–32 m long. Ventral microducts each
Zootaxa 2709 © 2010 Magnolia Press · 29
REVISION OF CRYPTOSTIGMA FERRIS
about 3.5 m wide, scattered evenly on venter. Tubular ducts absent. Clypeolabral shield 259–361 m wide.
Spiracular disc-pores each 4.5–5.3
m wide, with 5–7 (mostly 5 and 6) loculi, found in a small area around
spiracles. Multilocular disc-pores larger than spiracular disc-pores, each about 6.2
m wide, with 4–10
(mostly 7 & 8) loculi, numerous; present around vulva and in posterior abdominal segments. Antennae each
with 1–4 antennal segments, each very small, total length 38–70
m; with tips of fleshy setae mostly knobbed.
Interantennal setae about 3 pairs. Legs reduced, atrophied, with segments fused, total length 38–76
m. Tarsal
digitules similar, slender, usually spiniform, sometimes knobbed. Claw digitules, slender, knobbed. Claw
without a denticle. Spiracles larger than legs, located near margin; each spiracular peritreme 108–162
m
wide, anterior peritreme subequal to or slightly smaller than posterior peritreme.
Diagnosis. The adult female of C. inquilinum can be diagnosed by the following features: (i) dorsum with
numerous clusters of 4–10 small simple pores, (ii) dorsal setae flagellate, present in a single row around each
stigmatic sclerotization, absent elsewhere, (iii) dorsal microducts present, each bilocular, thick rimmed, (iv)
preopercular pores absent, (v) stigmatic sclerotization present, forming a sclerotized crescent, each with 2–3
stigmatic setae, (vi) each anal plate with 11–17 setae on dorsal surface, (vii) ventral tubular ducts absent, and
(viii) some fleshy setae on antennae always knobbed.
Remarks. The orbicular pores on the adult female C. inquilinum are often overlooked because they are
inconspicuous, and the borders of the membranous areas do not show signs of sclerotization as in other
species with orbicular pores. Hodgson (1994) and Qin and Gullan (1989) redescribed C. inquilinum, but
neither study mentions the presence of orbicular pores in this species. In the study by Qin and Gullan (1989),
the species is described as having no orbicular pores (as compound pores). The author believes that he has
examined the same material, or material from the same collections as Hodgson (1994) and Qin and Gullan
(1989), and that the orbicular pores were overlooked in these studies. The author only noticed the orbicular
pores of C. inquilinum after collecting fresh material in Mexico on Acacia paniculata, and later carefully
reexamining the type material listed above.
First-instar nymph (Fig. 10)
Unmounted material. Not available for study.
Mounted material. Elongate oval, 765–981
m long, 539–674 m wide (n=58).
Dorsum. Dorsal derm membranous, with segmentation delineated by membranous folds. Dorsal setae
slender, each 7–10
m long, present in 2 mid-dorsal longitudinal rows. Trilocular pore present on each side of
head near margin close to eye. Dorsal microducts each about 2
m wide, appearing bilocular, present
submarginally and in 4 submedian longitudinal rows. Simple pores each about 2
m wide, present
submarginally and in 2 submedian longitudinal rows. Anal plates triangular, each plate 92–107
m long, 30–
47
m wide; dorsal surface with 1 seta on anterior part of plate plus 3 apical setae; ventral surface with 1
fringe seta. Anal ring as in generic diagnosis. Eyespots present just above level of antennal scape.
Margin. Outline smooth. Marginal setae sharply spinose, each 15–20
m long; total number 60–66, with
about 8 anteriorly between eyes, and, on each side, 5 or 6 between each eye and anterior stigmatic setae, 7–9
between anterior and posterior stigmatic setae, and about 14–15 between each posterior stigmatic setae and
anal cleft. Stigmatic setae 1 per cleft, each bluntly spinose and 21–26
m long.
Ven te r. Ventral derm membranous. Antennae 6 segmented, total length 243–307
m. Ventral submedian
setae slender, with 1 pair on each of last 3 abdominal segments. Interantennal setae 1 pair. Submarginal setae
flagellate, with 7 pairs on each side of abdomen, a single seta between each anterior and posterior spiracles,
and 1 pair near apex of head. Ventral microducts each 2
m wide, about 6 on margin between eyes and
anterior stigmatic cleft, 3 submarginally between anterior and posterior stigmatic setae, and in pairs between
each inner and outer submarginal setae on abdomen. Spiracular disc-pores each about 4.5
m wide, with 5 or
6 loculi, each anterior stigmatic furrow with 7–8 pores, each posterior furrow with 7–9 pores. Spiracular
peritremes each 15–17
m wide. Clypeolabral shield 221–501 m wide. Legs well developed, with numerous
setae; trochanter + femur 132–153
m long, tibia + tarsus 153–170 m long, base of tarsi swollen in some
specimens studied (see asterisk (*) on Fig. 10); microctenidia present on apex of tibia; femur with 4–6 setae.
Tarsal digitules similar, slender. Claw slender, without a denticle; claw digitules slender, knobbed.
KONDO
30 · Zootaxa 2709 © 2010 Magnolia Press
Diagnosis. The first-instar nymph of C. inquilinum can be diagnosed by the following combination of
features: (i) dorsal setae present in 2 longitudinal rows of about 5 pairs, (ii) anterior and posterior stigmatic
clefts each with 1 stigmatic seta, (iii) with 7–9 marginal setae between anterior and posterior stigmatic setae,
(iv) antennae 6 segmented, (v) with 1 pair of ventral submedian setae on posterior 3 abdominal segments, (vi)
each anterior stigmatic furrow with 7–8 pores, each posterior furrow with 7–9 pores, and (vi) ventral
submarginal setae flagellate.
Remarks. The above description of the unmounted material was taken from the original description of
Akermes secretus Morrison (1922), a synonym of C. inquilinum. The ending of the specific epithetinquilina
is herein changed to “inquilinum” in order to match the neuter gender of the generic ending –stigma, in
accordance to the ICZN code (article 30.1.2.).
Host plants. Boraginaceae: Cordia alliodora. Fabaceae: Haematoxylon campechianum, Inga laurina,
Inga sp., Algarrobo (probably Prosopis juliflora). Moraceae: Ficus laevigata. Polygonaceae: Triplaris
surinamensis.
Associated Hymenoptera. Dolichoderinae: Azteca longiceps (Morrison, 1929); Formicinae:
Myrmelachista ambigua ramulorum Wheeler (Ferris, 1922); Myrmicinae: Crematogaster brevispinosa Mayr,
var. tumulifera (Newstead, 1920); Pseudomyrmecinae: Pseudomyrmex sp., allied to Ps. triplaridis Forel (as
Pseudomyrma) (Morrison, 1922).
Males. Unknown.
Distribution. Neotropical Region: Costa Rica, Cuba, Grenada, Guyana, Jamaica, Mexico, Panama,
Puerto Rico & Vieques Island, Trinidad & Tobago, U.S. Virgin Islands.
Cryptostigma jonmartini Kondo, sp. nov.
(Figs 1J & 11)
Material examined. Holotype. Adult
. GUATEMALA: Escuintla, Parque Auto Safari Chapín, 35 m asl,
(14°06’, 90°38’W), 14.xi.2003, coll. P.S. Ward, ex Enterolobium cyclocarpum (Fabaceae), 1 (1) (USNM).
Paratypes. Same data as holotype, 6 (6: 1 adult female + 4 third-instar nymphs + 1 first-instar nymph)
(USNM). MEXICO: Oaxaca, Temascal, 2.ii.1964, coll. J.H. Janzen, ex Astelia pterocarpa (Liliaceae), 5 (5: 4
adult female + 1 third-instar nymph), C.I.E 9420, B.M. 1968.27 (BMNH)
Adult female (Figs 1J & 11)
Unmounted material. Insects collected inside live hollow branches of host and tended by ants. The
following description is based on alcohol-preserved specimens. Third-instar nymphs yellow-cream; younger
adult females probably of a similar color. Mature adult females sclerotized, ferruginous, with black mottling
on dorsum; dorsum covered by a thin layer of wax. Mature adult females collected in Guatemala on
Enterolobium cyclocarpum become cylindrical in shape allowing them to fit perfectly into the hollow stems of
its host. The third-instar nymphs are flat in shape, and the insects only become cylindrical in the adult stage.
Mounted material. Body outline elongate, 3.3–6.0 mm long, 2.1–2.8 mm wide (n=6).
Dorsum. Derm membranous, sclerotized; with 3 subcircular orbicular pores, 1 on head region, and 1 on
each side between anterior and posterior stigmatic areas; each pore 160–330
m wide. Dorsum with
apodemes clearly visible, more or less evenly distributed on dorsum. Dorsal setae absent. Cribriform platelets
each 4.0–4.5
m wide, with 3–8 pores, each platelet square, hexagonal, triangular or irregular in shape,
sunken, scarce, present mostly around anal plates. Simple pores present, sunken, of 2 types, both evenly
scattered over dorsum: large pores each 6–7
m wide, thick; small pores each 3–4 m wide, scarce. Dorsal
microducts each 3–4
m wide, thick rimmed, sunken; evenly distributed on dorsum. Preopercular pores
absent. Sclerotic crescent present around anal plates. Anal plates together quadrate, with rounded angles, each
plate 183–203
m long, 70–88 m wide, anterolateral margin 110–125 m long, posterolateral margin 113–
135
m long; with 17–21 setae on dorsal surface; ventral subapical setae not visible due to sclerotization of
anal plates; plates located about 1/5–1/4 of body length from posterior margin. Anal ring with 10 setae. Eyes
absent.
Zootaxa 2709 © 2010 Magnolia Press · 31
REVISION OF CRYPTOSTIGMA FERRIS
Margin. Margin rugose. Marginal setae each 18–23 m long, sharply spinose, with well-developed bases,
arranged in a single irregular row, numerous, 30–41 between anterior and posterior stigmatic areas. Stigmatic
sclerotization present, forming a sclerotic crescent extending dorsad to spiracles. Stigmatic setae bluntly
spinose each 13–17
m long; with 1 in each cleft, often broken off.
Vent er. Derm membranous. Ventral setae slender, straight or slightly bent, each 13–23
m long, those near
margin shortest. Ventral microducts each 3–4
m wide, scattered evenly on venter. Tubular ducts absent.
Clypeolabral shield 285–375
m wide. Multilocular disc-pores each 4–6 m wide with 4–8 (mostly 6–8)
loculi, sunken; present medially on abdominal segments up to segment III, but more abundant around vulva
and in a marginal or submarginal band in abdominal region, forming a line of pores extending from vulvar
area alongside anal cleft to spiracular disc-pores of posterior spiracle. Spiracular disc-pores each 6–8
m
wide, with 5–8 (mostly 6–8) loculi, sunken, numerous, present in each spiracular furrow from laterad to each
spiracular peritreme to stigmatic cleft. Antennae reduced, 3 or 4 segmented, total length 105–130
m long;
with tips of fleshy setae commonly branching. Legs reduced, with most segments fused, atrophied; total
length 68–114
m. Tarsal digitules similar, knobbed. Claw digitules, slender, knobbed. Claw without a
denticle. Spiracles larger than legs, located near margin; each anterior peritreme 145–178
m wide, each
posterior peritreme 145–200
m wide.
Diagnosis. The adult female of C. jonmartini can be diagnosed by the following combination of
characters: (i) body shape elongate oval, (ii) dorsal setae absent, (iii) dorsum with 3 orbicular pores, (iv)
cribriform platelets present, (v) dorsal surface of anal plates with 17–21 setae, (vi) with 1 stigmatic seta per
cleft but often broken off, (vii) apodemes conspicuous on dorsal derm, (viii) many fleshy setae on antennae
branched, and (ix) presence of a marginal band of multilocular disc-pores around abdominal margin. C.
jonmartini appears morphologically closest to C. melissophilum. However, they can be separated by their
body shape and number of stigmatic setae (C. jonmartini being elongate oval with 1 stigmatic seta per cleft,
and C. melissophilum being oval with 3 stigmatic setae per cleft). In addition, C. jonmartini is associated with
tending ants whereas C. melissophilum is associated with bees. For a detailed list of differences, see Diagnosis
of C. melissophilum.
First-instar nymph (Fig. 12)
Unmounted material. Not available for study.
Mounted material. Oval, 1.1 mm long, 0.8 mm wide (n=1).
Dorsum. Dorsal derm membranous. Dorsal setae present in 2 mid-dorsal longitudinal rows, each seta 18
23
m long. Trilocular pore present on each side of head near margin anterior to eyes. Dorsal microducts 2 m
wide, appearing bilocular, present submarginally and in 2 submedian rows. Simple pores 3–5
m wide,
present submarginally and in 2 submedian longitudinal rows. Anal plates triangular, each plate 93–95
m
long, 25–28
m wide; dorsal surface with 1 seta on anterior part of each plate plus 3 apical setae; ventral
surface with 1 fringe seta. Anal ring as in generic diagnosis. Eyespots present on margin anterior to antennal
scape.
Margin. Outline smooth. Marginal setae sharply spinose, each 25–30
m long; total number 67, with 10
anteriorly between eyes, and, on each side, 7 or 8 between each eye and anterior stigmatic setae, 7 between
anterior and posterior stigmatic setae, and 14 between each group of posterior stigmatic setae and anal cleft.
Stigmatic setae each thick, bluntly spinose and 15–20
m long; with 1 per stigmatic cleft.
Ven te r. Ventral derm membranous. Antennae 6 segmented, total length 250–260
m. Ventral submedian
setae, slender, present in pairs on last 3 or 4 abdominal segments; also with a pair on thorax between meso-
and metathoracic legs. Interantennal setae 1 pair. Ventral submarginal setae slender, in 7 pairs on each side of
abdomen, outer submarginal setae longer than inner submarginal setae; and with 1 seta between each anterior
and posterior spiracles, and 1 pair near apex of head. Ventral microducts each about 2
m wide; with about 7
on submargin between each eye and anterior stigmatic cleft; 6 submarginally between each anterior and
posterior stigmatic setae, and about 2 near inner submarginal setae on abdomen; only 1 present next to
penultimate inner submarginal setae, but absent from pair near anal cleft and from pair just below posterior
spiracular furrow on each side. Spiracular disc-pores each about 4
m wide, with 5–7 (mostly 5) loculi;
stigmatic furrows each with 7–8 pores. Spiracular peritremes each 10–13
m wide. Clypeolabral shield 175
KONDO
32 · Zootaxa 2709 © 2010 Magnolia Press
m wide. Legs well developed; trochanter + femur 123–133 m long, tibia + tarsus 143–155 m long. Femur
with 2 closely-paired setae (see arrow and circlet on Fig. 6). All tarsal digitules similar, slender. Claw without
a denticle; claw digitules slender, knobbed, 1 slightly thicker than other.
FIGURE 11. Cryptostigma jonmartini Kondo, adult female.
Zootaxa 2709 © 2010 Magnolia Press · 33
REVISION OF CRYPTOSTIGMA FERRIS
FIGURE 12. Cryptostigma jonmartini Kondo, first-instar nymph.
Diagnosis. The first-instar nymph of C. jonmartini can be diagnosed by the following features: (i) dorsal
setae present in 2 longitudinal rows of about 5 pairs, (ii) with 1 stigmatic seta in each anterior and posterior
stigmatic cleft, (iii) with 7 marginal setae between anterior and posterior stigmatic setae, (iv) antennae 6
KONDO
34 · Zootaxa 2709 © 2010 Magnolia Press
segmented, (v) with 4 or 5 pairs of ventral submedian setae, including 1 pair present on thorax between meso-
and metathoracic region, (vi) femur with 2 closely-paired setae, and (vii) spiracular furrows each with 7–8
pores.
Host plants. Fabaceae: Enterolobium cyclocarpum. Liliaceae: Astelia pterocarpa.
Associated Hymenoptera. Formicidae: Pseudomyrmecinae: Pseudomyrmex sp.
Etymology. The species is named after Dr. Jon Martin, curator of the suborder Sternorrhyncha
(Hemiptera) at the Natural History Museum, London. Dr. Martin is an authority of the whiteflies
(Aleyrodidae), and is also a taxonomic specialist of aphids (Aphididae), jumping plant lice (Psyllidae) and
scale insects (Coccoidea) and has collected many scale insect species, including some described here.
Males. Unknown.
Distribution. Neotropical Region: Guatemala, Mexico.
Cryptostigma longinoi Kondo, sp. nov.
(Figs 1G & 13)
Material examined. Holotype. Adult
. COSTA RICA: 1 (1), Provincia Puntarenas, Ojo de Agua, 800m asl,
10°16’N, 84°50’W, 5.vii.1991, coll. J. Longino #2956, ex nest of Azteca longiceps in Triplaris
melaenodendron, (USNM). Paratypes. Same data as holotype: 7 (7: 2 adult
+ 4 third-instar nymphs + 1
first-instar nymph) (USNM); Puntarenas, rainforest 19 km S. Ciudad Neily, 20 m asl, 8°29’N, 82°58’W,
26.iii.1990, coll. P. S. Ward, # 10650, ex live branch with Azteca 2 (2 third-instar nymphs), (USNM);
Puntarenas, Reserva Biológica Carara, 9°47’S, 84°35’W, 400m asl, 15.ix.1988, coll. J. Longino (# 2196), ex
Cecropia insignis with Azteca xanthochroa 2 (2) (USNM, BME); Provincia Guanacaste, Parque Nacional
Palo Verde, -.i.2001, coll. V. Carmona, A. Carmona and T. Valdieso, ex Cecropia peltata, in hollow stems,
tended by Azteca sp. 2 (2) (AUCC).
Other material studied. SURINAM: Paramaribo, 9.viii.1932, coll. G. Bunzli, No, -.VIII.26, within
Cecropia, attended by ant, 3 (4: 1 adult female + 1 third-instar nymph + 2 first-instar nymphs) (BMNH).
Adult female (Figs 1G & 13)
Unmounted material. Alcohol-preserved specimens (Fig. 1G) subcircular; young adult female yellow
cream, dorsum speckled with tiny dark spots corresponding to sclerotized pores on slide-mounted specimens;
anal plates and surrounding area dark; stigmatic clefts also dark, with a white soft wax. Color of matured adult
female yellow cream mixed with reddish brown, interrupted by transverse dark rows. Body becoming
sclerotized around margins.
Mounted material. Body outline oval to broadly oval, margins smooth or irregular, undulated, 2.5–5.4
mm long, 2.3–4.2 mm wide (n=5).
Dorsum. Derm membranous, with numerous circular or irregularly-shaped sclerotic pores, each pore 13–
43
m wide, of 2 types: i) sclerotic pores with a wide central locule (most pores on Fig. 12) present around
anal plates and mid-dorsal area, and ii) sclerotic pores with a small central locule present throughout dorsum;
with an area with few or no pores around anal plates. Dorsal setae knobbed or bluntly spinose, but with some
sharply spinose, each seta 10–15
m long, with well-developed base, most on a small sclerotic plate, 3–15 m
wide; base of each seta usually about 1/3 length of seta; present evenly on dorsum. Preopercular pores absent.
Dorsal microducts each 2–3
m wide, numerous, with a very long outer ductule; inner ductule about as long
as or longer than outer ductule; scattered on dorsum. Sclerotic area present around anal plates; derm sunken
around anal plates and usually devoid of sclerotic pores; older specimens with a C-shaped (inverted C-shape
on left side) sclerotization at base of each anal plate, which in very old specimens, becomes fused to
sclerotized area around anal plates (see enlargement on right side of Figure 12). Anal plates together quadrate
to pyriform, with rounded angles; each plate 200–253
m long, 80–105 m wide, anterolateral margin 138–
198
m wide, posterolateral margin 112–138 m wide; with 3–6 setae on dorsal surface of each plate; plates
located about 1/3–1/4 of body length from posterior margin. Anal lobes closely adpressed along entire length,
and often becoming entirely fused and leaving no signs of an anal cleft. Anal ring with 10 setae. Eyes absent.
Zootaxa 2709 © 2010 Magnolia Press · 35
REVISION OF CRYPTOSTIGMA FERRIS
FIGURE 13. Cryptostigma longinoi Kondo, adult female.
KONDO
36 · Zootaxa 2709 © 2010 Magnolia Press
Margin. Marginal setae not differentiated. Stigmatic sclerotization rather small; stigmatic setae not
detected.
Ve nt er. Derm membranous. Ventral setae slender, straight or slightly bent, each 13–38
m long, longest
around vulvar region and on abdominal segments. Interantennal setae 2 or 3 pairs. Ventral microducts each
about 2.0–2.5
m wide, scattered evenly on venter. Tubular ducts absent. Clypeolabral shield 325–420 m
wide. Multilocular disc-pores each 5–8
m wide, with 3–8 (mostly 5–7) loculi, abundant around vulva and
present on all abdominal segments plus metathoracic segment; some specimens also with 1 or a few pores
laterad to mouthparts. Simple pores present, each 4.5–7.5
m wide, located on thorax, especially on pro- and
metathoracic segments, often with a few also around meso- and metathoracic legs, and 1 or 2 elsewhere on
thoracic region; rarely only 1 present on entire venter, near mouthparts (see Remarks). Spiracular disc-pores
each 5–6
m wide, each with 3–8 (mostly 5 & 6) loculi; with a line of pores extending anteriorly towards
mesothoracic legs. Antennae small, with 1–3 fused segments, total length 33–50
m. Legs greatly reduced,
represented by a rudimentary claw, several setae, and associated pores, total length 13–25
m. Spiracles much
larger than legs, located near body margin just below stigmatic sclerotization; anterior spiracular peritremes
each 105–170
m wide, posterior peritremes each 125–170 m wide.
Diagnosis. The adult female of C. longinoi can be diagnosed by the following features: (i) dorsum with
numerous sclerotic pores, (ii) dorsal setae present, knobbed or bluntly spinose, some sharply spinose, (iii)
preopercular pores absent, (iv) each anal plate with 3–6 setae on dorsal surface, (v) ventral tubular ducts
absent, and (viii) simple pores present on venter. This is the only known species with simple pores on the
venter.
First-instar nymph (Not illustrated).
Unmounted material. Not available for study.
Mounted material. Elongate oval, 1.2 mm long, 0.8 mm wide (n=1).
Dorsum. Dorsal derm membranous. Dorsal setae slender, each 5–13
m long, present in 2 submedian
parallel rows. Trilocular pore present on each side of head near margin close to eye. Dorsal microducts each
about 2
m wide, with orifice appearing bilocular, apparently present submarginally around body and in 2
submedian longitudinal rows. Simple pores each 2.5–3.5
m wide, distribution similar to dorsal microducts.
Anal plates with shingled surface texture, together pyriform, each plate 98
m long, 45 m wide, anterolateral
margin 78
m long, posterolateral margin 55 m long, dorsal surface with 1 seta on anterior part of plate plus
about 4 apical setae; ventral surface with 1 fringe seta. Anal ring as in generic diagnosis.
Margin. Outline smooth. Marginal setae sharply spinose, each 20–28
m long, total number 75, with
about 10 anteriorly between eyes, and on each side, about 8 between each eye and anterior stigmatic setae, 8
between each anterior and posterior stigmatic setae, and 13–15 between posterior stigmatic setae and anal
cleft. Stigmatic setae sharply spinose, some with rounded tips, all subequal in length or median seta longest,
each seta 13–20
m long; with 3 per stigmatic cleft.
Venter. Ventral derm membranous, with microtrichia present on mid-areas of abdominal segments.
Antennae 5 segmented, total length 290
m. Interantennal setae 1 pair. Ventral submedian setae: 1 pair on 3
last abdominal segments. Ventral submarginal setae slender, outer submarginal setae 7 pairs, inner
submarginal setae 6 pairs; also 2 setae between each anterior and posterior spiracles, and 1 pair near apex of
head. Ventral microducts each about 2
m wide, present in a single row around body margin. Spiracular disc-
pores each 4
m wide, with 4–7 (mostly 5) loculi; with with 6–8 pores in each anterior stigmatic furrow and
9–10 pores in each posterior furrow. Spiracular peritremes each 17.5
m wide. Clypeolabral shield 158 m
wide. Legs well developed; trochanter + femur 148–155
m long, tibia + tarsus 170–175 m long; femur with
6–8 setae. Prothoracic tarsal digitules dissimilar, 1 knobbed and 1 slender. Meso- and metathoracic tarsal
digitules similar, knobbed. Claw without a denticle; claw digitules slender, knobbed, 1 slightly thicker than
the other.
Diagnosis. The first-instar nymph of C. longinoi can be diagnosed by the following combination of
features: (i) dorsal setae present in 2 longitudinal rows, (ii) with 3 stigmatic setae in each stigmatic cleft, (iii)
with 8 marginal setae between anterior and posterior stigmatic setae, (iv) antennae 5 segmented, (v) 1 pair of
ventral submedian setae on last 3 abdominal segments, (vi) anterior stigmatic furrows each with 6–8 pores,
Zootaxa 2709 © 2010 Magnolia Press · 37
REVISION OF CRYPTOSTIGMA FERRIS
posterior stigmatic furrows each with 9–10 pores, and (vi) femur with 6–8 setae. The preceding description of
the first-instar nymph is based on a fully matured first-instar nymph from Costa Rica, close to molting and in
poor condition.
Variation. All adult specimens from Costa Rica have easily discernible thick-rimmed ventral disc pores,
but the only specimen available from Surinam has only one simple pore near the mouthparts; and the two
first-instar nymphs from Surinam are in poor condition but have only 4 or 5 setae in each femur in contrast to
6–8 setae observed in the type material. Despite the differences observed in the Surinam material, the
specimens appear to share sufficient features with the specimens from Costa Rica and are considered here to
be conspecific.
Host plants. Cecropiaceae: Cecropia insignis, Cecropia peltata; Polygonaceae: Triplaris
melaenodendron.
Associated Hymenoptera. Formicidae: Dolichoderinae: Azteca longiceps, Azteca xanthochroa, Azteca
sp.
Males. Unknown.
Etymology. The species is name after Dr. John T. Longino who collected the holotype. Dr. Longino is a
world authority on neotropical myrmecology.
Distribution. Neotropical Region: Costa Rica, Surinam.
Cryptostigma melissophilum Kondo, sp. nov.
(Figs 1A, 14 & 15)
Material examined. Holotype. Adult
. BRAZIL: Amazonas, Rio Negro, Tapurucuara Mirim, 0º25’17”S,
66º24”22”W, 3.vii.1999, coll. João M.F. Camargo and Silvia R.M. Pedro, ex Campsiandra angustifolia, inside
nest of Schwarzula coccidophila, series number 741c (Apidae: Meliponini), AL-051-2001, 1 (1) (MZSP).
Paratypes. Same data as holotype 2 (3: 1 adult
+ 2 first-instar nymphs) (MZSP); 29 (RPSP) in alcohol; 7 (7:
4 adult
+ 3 first-instar nymphs) (USNM); same data as holotype but 2.vii.1999, 4 (4) (USNM), 75 (RPSP)
in alcohol.
Adult female (Figs 1A & 14)
Unmounted material. Young adult female subcircular, flattish, constricted at stigmatic clefts; entire body
covered with small grayish white waxy flakes; color of body peach pink after removal of wax, with orbicular
pores visible as tiny red-brown spots, and with a somewhat powdery wax present just around each orbicular
pore; anal plates and circular area around plates reddish to purplish brown; anal cleft only just visible; area
around spiracular peritremes conspicuously marked by white spots made of a wax secreted by spiracular disc-
pores. Mature adult female highly convex, oval to elongate oval, becoming irregular in shape when crowded,
dorsum with a somewhat granulose texture, body constricted at the stigmatic clefts; color of insect peach pink;
dorsum with 6–7 yellowish-white transverse intersegmental lines separated from each other at regular
intervals; transverse lines interrupted by a median longitudinal groove and 1 or 2 additional grooves running
from thoracic region near head posteriorly towards area just anterior or laterad to anal plates. Orbicular pores
present close to body margin, represented by 3 tiny circular reddish-brown spots: 1 on head region, and 1
between stigmatic areas on each side of body. Area around spiracular peritreme marked by snow-white wax.
Different growth stages usually found within each colony, with first- and second-instar nymphs appearing
yellowish.
Mounted material. Body outline oval, constricted in areas of stigmatic clefts; 6.0–11.0 mm long, 5.2–8.5
mm wide (n=10).
KONDO
38 · Zootaxa 2709 © 2010 Magnolia Press
FIGURE 14. Cryptostigma melissophilum Kondo, adult female.
Dorsum. Derm membranous; with 3 subcircular orbicular pores: 1 on head region, and 1 on each side
between anterior and posterior stigmatic areas; each pore 162–329
m wide. Dorsum with apodemes clearly
visible, evenly distributed on dorsum; most conspicuous on submarginal area. Dorsal setae absent. Cribriform
Zootaxa 2709 © 2010 Magnolia Press · 39
REVISION OF CRYPTOSTIGMA FERRIS
platelets each 7–11 m wide, with 4–8 pores, each platelet square, hexagonal or irregular in shape. Simple
pores of 2 types: large simple pores each 7–11
m wide, thick; numerous, scattered evenly on dorsum; small
simple pores each 2–3
m wide; numerous, evenly distributed on dorsum. Dorsal microducts unilocular, each
about 3–4
m wide, thick rimmed, sunken; evenly distributed on dorsum. Preopercular pores absent. Sclerotic
crescent present around anal plates. Anal plates together quadrate, with rounded angles, each plate 194–232
m long, 102–113 m wide, with about 16 setae on dorsal surface, plus about 7 ventral subapical setae on
each plate; plates located about 1/5–1/4 of body length from posterior margin. Anal ring with 10 setae. Eyes
absent.
Margin. Margin rugose. Marginal setae present, each 15–22
m long and sharply spinose, with well-
developed bases; arranged in a single irregular row, numerous, about 32–45 between anterior and posterior
stigmatic areas. Stigmatic clefts very deep, stigmatic sclerotization extending to dorsad to spiracles. Stigmatic
setae each subequal in length (15–21
m long) and bluntly spinose, with 3 in each cleft but often broken off.
Venter. Derm membranous. Ventral setae slender, straight or slightly bent, each 7–15
m long, abundant
on abdominal segments and toward body margin. Ventral microducts each about 3–4
m wide, scattered on
venter, numerous around labium. Tubular ducts absent. Clypeolabral shield 361–426
m wide. Multilocular
disc-pores each 4–6
m wide with 5–9 (mostly with 8) loculi, sunken; abundant around vulva and in a
submarginal band in abdominal region, forming a line of pores extending from vulva alongside anal cleft to
spiracular disc-pores of posterior spiracle; pores also present on mid-venter on all abdominal segments.
Spiracular disc-pores each 4–6
m wide with 5–8 (mostly 8) loculi, sunken; numerous, found within a
spiracular furrow extending from laterad to each spiracular peritreme to stigmatic cleft. Antennae reduced,
with 3–8 antennal segments, total length 94–117
m; tips of fleshy setae commonly bifurcating. Legs reduced,
atrophied; with most segments fused, total length 47–149
m. Prothoracic tarsal digitules dissimilar, 1
slender, 1 knobbed. Meso- and metathoracic tarsal digitules similar, knobbed. Claw digitules, slender,
knobbed. Claw without a denticle. Spiracles larger than legs, located near margin. Anterior spiracular
peritremes each 216–259
m wide, posterior peritremes each 243–297 m wide.
Diagnosis. The adult female of C. melissophilum can be diagnosed by the following features: (i) body
shape oval, (ii) dorsal setae absent, (iii) dorsum with 3 orbicular pores, (iv) cribriform platelets present, (v)
anal plates with about 16 setae on dorsal surface, (vi) stigmatic setae totalling 3 per cleft, but often broken off,
(vii) apodemes conspicuous on dorsal derm marginally and submarginally, (viii) some fleshy setae on
antennae always branched, and (ix) presence of a marginal band of multilocular disc-pores around body
margin on abdomen. C. melissophilum appears most similar in morphology to C. jonmartini. Besides the
morphological differences between the two species given in the key to adult female Cryptostigma, C.
melissophilum and C. jonmartini can be separated by the following character states: (i) the thickness of the
waxy layer covering the dorsum of live specimens (rather thick, about 1.5 mm or more in C. melissophilum;
thin, less than 1 mm in C. jonmartini); and (ii) the width of posterior spiracular peritremes (each 243–297
m
wide in C. melissophilum; 145–200
m wide in C. jonmartini). In addition, C. jonmartini is associated with
tending ants whereas C. melissophilum is associated with bees.
Remarks. Cryptostigma melissophilum, as the name suggests, is closely associated with stingless bees.
Camargo and Pedro (2002a) discovered and reported in detail the bee-coccid association and identified the
bee as Schwarzula sp. (now Schwarzula coccidophila Camargo & Pedro, 2002b), family Apidae: tribe
Meliponini. The stingless bees obtain honeydew for food and also collect wax from the waxy test to build
their nest (Camargo and Pedro, 2002a). C. melissophilum has a rather thick waxy layer which is constantly
harvested by the stingless bees which rub the wax between their basitarsi to form pellets, which they then
transport in their mandibles to their nest where it is stored as wax deposits or used in the construction or repair
of their nest (Camargo & Pedro 2002a). The thicker wax layer in C. melissophilum may therefore be an
adaptation to this mutualistic relationship with the stingless bees. The wax of ant-tended coccid species does
not seem to be collected or used by tending ants, which appear to be only interested in the honeydew.
Symbiotic relationships in the Apidae are rare, with only about 3 reports known (J.M.F. Camargo, personal
communication).
First-instar nymph (Fig. 15)
KONDO
40 · Zootaxa 2709 © 2010 Magnolia Press
FIGURE 15. Cryptostigma melissophilum Kondo, first-instar nymph.
Unmounted material. Not available for study.
Mounted material. Elongate oval, 1024–1073
m long, 733–782 m wide (n=5).
Zootaxa 2709 © 2010 Magnolia Press · 41
REVISION OF CRYPTOSTIGMA FERRIS
Dorsum. Dorsal derm membranous, with segmentation delineated by membranous folds. Dorsal setae
each 13–16
m long, present in 2 mid-dorsal longitudinal rows. Trilocular pore present on each side of head
near margin close to eye. Dorsal microducts each 2–4
m wide, appearing bilocular, present submarginally
and in 4 submedian rows. Simple pores each 3–4
m wide, present submarginally and in 2 submedian
longitudinal rows. Anal plates triangular, each plate 100–104
m long, 11–32 m wide; dorsal surface with 1
seta on anterior part of plate plus 3 apical setae; ventral surface with 1 fringe seta. Anal ring with 6 setae plus
an irregular row of translucent wax pores. Eyespots present just above level of antennal scape.
Margin. Outline smooth. Marginal setae sharply spinose, each 26–36
m long; total number 73–82, with
10–11 anteriorly between eyes, 7–11 between each eye and anterior stigmatic seta, 9–11 between each
anterior and posterior stigmatic setae, and 13–16 between each posterior stigmatic seta and anal cleft.
Stigmatic setae thick, bluntly spinose, each 22–30
m long; with 1 per stigmatic cleft.
Ven te r. Ventral derm membranous. Antennae 6 segmented, total length 286–318
m. Ventral submedian
setae, slender, present in pairs on last 3 abdominal segments, plus a pair on thoracic region between meso- and
metathoracic legs. Interantennal setae 1 pair. Ventral submarginal setae slender, with outer submarginal setae
much longer than inner submarginal setae, present in 7 pairs on abdomen on each side, plus a single seta
between each anterior and posterior spiracles, and 1 pair near apex of head. Ventral microducts each about 2
m wide, with about 11 on submargin between each eye and anterior stigmatic cleft; and on each side, 7
submarginally between each anterior and posterior stigmatic setae, and about 4 or 5 near pairs of submarginal
setae on abdomen, but absent from near anal cleft. Spiracular disc-pores each 4–6
m wide with 5–8 (mostly
7) loculi; anterior stigmatic furrows each with 8–11 pores, posterior furrows each with 9–12 pores. Spiracular
peritremes each 18–20
m wide. Clypeolabral shield 226–232 m wide. Legs well developed, with numerous
setae; trochanter + femur 128–150
m long, tibia + tarsus 139–160 m long; microctenidia present on apex of
tibia. Tarsal digitules similar, slender. Claw without a denticle; claw digitules slender, knobbed.
Diagnosis. The first-instar nymph of C. melissophilum can be diagnosed by the following combination of
features: (i) dorsal setae present in 2 longitudinal rows of about 5 pairs, (ii) with 1 stigmatic seta in each
anterior and posterior stigmatic cleft, (iii) presence of 7 marginal setae between anterior and posterior
stigmatic setae, (iv) antennae 6 segmented, (v) ventral submedian setae 4 pairs: 1 pair on last 3 abdominal
segments, plus 1 isolated pair between meso- and metathoracic legs (vi) anterior stigmatic furrows each with
8–11 pores, posterior stigmatic furrows each with 9–12 pores, (vii) presence of 3 or 4 microducts between
each pair of submarginal setae on abdomen and (viii) femur with 4 setae.
Host plants. Fabaceae: Campsiandra angustifolia.
Associated Hymenoptera. Apidae: Meliponini: Schwarzula coccidophila Camargo & Pedro, 2002b.
Males. Unknown.
Etymology. The specific epithet: “melissophilum” is composed of the Greek wordsmelissa” meaning
bee and “philum” which is a neutral adjective meaning loving. Named after its close association with stingless
bees.
Distribution. Neotropical Region: Brazil.
Cryptostigma mexicanum Kondo, sp. nov.
(Fig. 16)
Material examined. Holotype. Adult
. MEXICO: Intercepted at Dallas, Fort Worth, 31.vii.1978, coll. G.M.
Stamey, in wood stick of crate (from Mexico), 1 (3: holotype clearly indicated on label; smallest of three
specimens, located centermost, dimensions: 2.7 mm wide, 3.0 mm long) (USNM). Paratypes. Same data as
holotype 2 (9: 3 adult
+ 6 immatures) (USNM); MEXICO: Jalisco, San Antonio, 4.ix.1976, coll. D.
Johnston, #8535 (77–31), ex hollow pit of twig of wood crate, 2 (4: 2 adult
+ 1 third-instar + 1 immature
[2
nd
instar] ) (USNM); MEXICO: Intercepted at New York, date not given, coll. C.E. Stegmaier, ex
Sambucus wood from Mexico, 1 (3) (USNM).
Adult female (Fig. 16)
Unmounted material. Not available during present study.
KONDO
42 · Zootaxa 2709 © 2010 Magnolia Press
FIGURE 16. Cryptostigma mexicanum Kondo, adult female.
Mounted material. Body outline oval to elongate oval, 2.5–3.3 mm long, 2.1–3.1 mm wide (n=8).
Dorsum. Derm membranous. Dorsal setae slender, with a straight or bent apex, each 11–18
m long;
present evenly on dorsum. Simple pores circular to subcircular; variable in size, large pores 3–4
m wide,
Zootaxa 2709 © 2010 Magnolia Press · 43
REVISION OF CRYPTOSTIGMA FERRIS
small pores about 2 m wide, probably minute microducts; numerous, scattered evenly on dorsum. Dorsal
microducts each 3–4
m wide, thick-rimmed, numerous, evenly distributed on dorsum. Preopercular pores
present, each 5.5–10
m wide; few, totalling 10–30, located in a small area around anterolateral margin of
anal plates. Sclerotic crescent present around anal plates. Anal plates together slightly pyriform, with rounded
angles, each plate 183–243
m long, 86–97 m wide, anterolateral margin 162–189 m long, posterolateral
margin 108–113
m long; with about 10 setae on dorsal surface, plus about 3 ventral subapical setae on each
plate; plates located about 1/5 of body length from posterior margin. Anal ring with 10 setae. Eyes absent.
Margin. Margin smooth. Marginal setae present, sharply spinose, short, each 8.9–16.0
m long, arranged
in 1 or 2 rows, mostly in 1 row, with 16–36 setae between anterior and posterior stigmatic areas. Stigmatic
clefts very deep; stigmatic sclerotization extending from margin to dorsad to spiracles. Stigmatic setae not
detected.
Vent er. Derm membranous. Ventral setae slender, straight or slightly bent, each 15–36
m long, those on
abdominal segments usually longer. Ventral microducts each about 3
m wide, appearing bilocular, scattered
evenly on venter. Tubular ducts absent. Clypeolabral shield 329–372
m wide. Multilocular disc-pores fairly
large, each 6–8
m wide, with 4–8 loculi, present around vulvar area, and in transverse bands in mid-ventral
areas of all abdominal segments and extending onto meso- and metathoracic region; also with small
subgroups of pores present next to each abdominal pore band. Spiracular disc-pores each 4–6
m wide with
4–7 loculi, present around spiracles and reaching body margin, with a line of pores connecting from each
spiracle to each prothoracic and mesothoracic leg respectively. Antennae very small, 2–4 segmented, total
length 26–53
m; with fleshy setae often bifurcated. Interantennal setae about 4 pairs. Legs greatly reduced,
each with about 4 segments; each segment mostly quadrate, rarely rounded; total length 30–104
m. Claw
present, without a denticle. Spiracles much larger than legs, located near margin; anterior spiracular
peritremes each 124–168
m wide, posterior peritremes each 124–177 m wide.
Diagnosis. The adult female of C. mexicanum can be diagnosed by the following features: (i) dorsum
without sclerotic pores, (ii) dorsal setae present, slender, scattered throughout dorsum, (iii) dorsum with 10–
30 preopercular pores, (iv) multilocular disc-pores present around pro- and mesothoracic legs, (v) leg
segments usually characteristically square or rectangular in shape (vi) presence of about 10 setae on surface of
each anal plate, and (vii) ventral tubular ducts absent.
Remarks. First-instar nymphs were not available during present study. Cryptostigma mexicanum has
been collected only from twigs of wooden crates originating from Mexico at USA quarantine facilities at New
York and Texas.
Host plants. Caprifoliaceae: Sambucus sp.
Associated Hymenoptera. None reported.
Males. One second-instar male was present among the material studied.
Etymology. The species is named after its country of origin: Mexico.
Distribution. Neotropical Region: Mexico.
Cryptostigma philwardi Kondo, sp. nov.
(Fig. 17)
Material examined. Holotype. Adult
. COLOMBIA: Magdalena, Ciénaga, 11°00’N, 74°15’W,
15.viii.1985, coll. P.S. Ward (#7932), ex colony of Pseudomyrmex sp. cf. fortis, on live tree of Avicennia
germinans, 1 (1) (USNM). Paratype. Same data as holotype 1 (1) (USNM).
Adult female (Fig. 17)
Unmounted material. Not available during present study.
Mounted material. Body outline oval, 3.0–4.7 mm long, 2.2–3.0 mm wide (n=2).
KONDO
44 · Zootaxa 2709 © 2010 Magnolia Press
FIGURE 17. Cryptostigma philwardi Kondo, adult female.
Dorsum. Derm membranous, with elevated ridges arranged in a cellular, tessellated pattern, transverse
ridges well-developed on thorax and abdomen. Orbicular pores present, each pore 125–305
m wide, with 1
Zootaxa 2709 © 2010 Magnolia Press · 45
REVISION OF CRYPTOSTIGMA FERRIS
or 3 on head region and 2 or 5 on thoracic region, largest on head. Dorsal setae absent, except for a group of
25–40 flagellate setae present next to each stigmatic sclerotization on a thin membranous area, each 15–55
m
long. Simple pores present, of 3 types: large simple pores each 5–6
m wide, scattered evenly on dorsum;
small simple pores each 2–3
m wide, evenly distributed on dorsum, and a second type of small simple pores
each 3–4
m wide, arranged in groups of 2–15 pores scattered throughout dorsum but most abundant near
anal plates. Dorsal microducts bilocular, thick rimmed, each 4–5
m wide, evenly distributed on dorsum.
Preopercular pores absent. Sclerotic crescent present around anal plates. Anal plates together quadrate, with
rounded angles, each plate 215–220
m long, 88–100 m wide, anterolateral margin 125–175 µm long,
posterolateral margin 130–135 µm long; with 12–27 setae on dorsal surface; ventral subapical setae not
visible due to sclerotization of anal plates; plates located about 1/4 of body length from posterior margin. Anal
ring with 10 setae. Eyes absent.
Margin. Margin rugose, smooth or undulating. Marginal setae each 13–18
m long and sharply spinose,
with well-developed bases; arranged in a single irregular row, with 40–60 between anterior and posterior
stigmatic areas. Stigmatic clefts very deep, stigmatic sclerotization C-shaped, closely associated with
spiracles. Stigmatic setae bluntly spinose with swollen bases, rarely sharply spinose, each cleft with 2 or 3
setae 15–18
m long but often broken off.
Ve nt er. Derm membranous. Ventral setae slender, straight or slightly bent, each 18–35
m long. Ventral
microducts each 3–4
m wide, scattered evenly on venter. Tubular ducts absent. Clypeolabral shield 285 m
wide. Multilocular disc-pores each 6–7
m wide, with 5–8 (mostly 8) loculi, abundant around vulvar area,
extending onto last 2 abdominal segments. Spiracular disc-pores each 5–6
m wide, with 5–7 (mostly 6–7)
loculi, found closely associated with each spiracle. Antennae reduced, with 1 or 2 antennal segments, total
length 53–65
m. Legs reduced, atrophied; with most segments fused, total length 33–60 m. Tarsal digitules
similar, long, setose. Claw digitules, slender, knobbed. Claws well developed, without a denticle. Spiracles
larger than legs, located near margin; anterior spiracular peritremes each 113–125
m wide, posterior
peritremes each 123–130
m wide.
Diagnosis. The adult female of C. philwardi can be diagnosed by the following features: (i) derm with
elevated ridges arranged in a cellular or tessellated pattern, (ii) sclerotic pores absent, (iii) dorsum with
numerous clusters of 2–15 small simple pores, (iv) dorsal setae absent, except for a group of 25–40 flagellate
setae present next to each stigmatic sclerotization on a thin membranous area, (v) orbicular pores present, with
1 or 3 on head region and 2 or 5 on thoracic region, (vi) preopercular pores absent, (vii) stigmatic
sclerotization forming a sclerotized crescent, each with 2 or 3 stigmatic setae, (viii) each anal plate with 12–27
setae on dorsal surface, and (ix) ventral tubular ducts absent. C. philwardi is the only known species in the
genus with a cellular or tessellated pattern on the dorsum.
Remarks. First-instar nymphs were not available during present study.
Host plants. Verbenaceae: Avicennia germinans.
Associated Hymenoptera. Formicidae: Pseudomyrmecinae: Pseudomyrmex sp. cf. fortis.
Males. Unknown.
Etymology. The species is named after Dr. Philip S. Ward, its collector and world authority of ant
systematics, particularly those ants in the subfamily Pseudomyrmecinae .
Distribution. Neotropical Region: Colombia.
Cryptostigma reticulolaminae Morrison
(Figs 1D, 18 & 19)
Cryptostigma reticulolaminae Morrison; Morrison 1929: 51.
Material examined. Holotype. Adult . PANAMA (AS CANAL ZONE): Frijoles, 28.iii.1923, coll. W.M.
Wheeler (#232), ex Cordia alliodora, 1 (1) (USNM). Paratypes. Same data as holotype, but #186, 3 (3)
(USNM).
KONDO
46 · Zootaxa 2709 © 2010 Magnolia Press
FIGURE 18. Cryptostigma reticulolaminae Morrison, adult female.
Zootaxa 2709 © 2010 Magnolia Press · 47
REVISION OF CRYPTOSTIGMA FERRIS
FIGURE 19. Cryptostigma reticulolaminae Morrison, first-instar nymph.
KONDO
48 · Zootaxa 2709 © 2010 Magnolia Press
Other material examined. COSTA RICA: Guanacaste Prov., Palo Verde, 15.vii.1979, coll. R.F.D. (II-2-
80), ex stem of Triplaris Americana, tended by Pseudomyrmex triplaridis, 6 (7) (USNM); MEXICO:
Chamela, -.ix.2005, coll. D. Gordon and R. Dirzo, ex Cordia alliodora, inside hollow stem and tended by
ants, 3 (3: 2 adult females + 1 third-instar nymph) (BME); PANAMA (AS CANAL ZONE): Barro Colorado
Island, 2.viii.1924, coll. W.M. Wheeler (#832), 2 (2) (USNM); Ancon, 18.iii.1923, coll. W.M. Wheeler
(#186), ex Cordia alliodora, 2 (20: 6 first-instar nymphs + 14 embrionic larvae) (USNM).
Adult female (Figs 1D & 18)
Unmounted material. “Specimens varying in size and shape, average length 5 mm., width 4.5 mm.,
height 2.5 mm., deeply invaginated beneath in the abdominal region; color of alcoholic specimens dark
reddish brown with blackish infusion along margin and anterior to anal plates; body surface smooth, rather
shining, bearing numerous tiny pits corresponding in position to sclerotic plates of derm” (Morrison, 1929).
Mounted material. Body outline oval to elongate oval, 2.8–6.0 mm long, 2.2–5.0 mm wide (n=9).
Dorsum. Derm membranous, with numerous subcircular sclerotic pores arranged in a reticulate pattern,
each pore 30–107
m wide. Dorsal setae sharply to bluntly spinose, with a swollen or spatulate apex, each 11–
22
m long, scattered evenly on dorsum between sclerotic pores. Simple pores not detected. Dorsal
microducts heavily sclerotized, each 4–5
m wide, outer ductule deep, broadened near duct opening; outer
ductule appearing longer than inner ductule; numerous, evenly distributed on dorsum. Preopercular pores
absent. Narrow sclerotic crescent present around anal plates. Anal plates together quadrate, with rounded
angles, each plate 149–181
m long, 64–90 m wide, anterolateral margin 75–113 m long, posterolateral
margin 117–134
m long; with about 4 setae on dorsal surface, plus about 3 ventral subapical setae on each
plate; plates located about 1/4 of body length from posterior margin. Anal ring with 10 setae. Eyes absent.
Margin. Margins becoming heavily sclerotized at maturity, with sclerotic pores around body margin.
Marginal setae not differentiated from dorsal setae. Stigmatic clefts very deep; stigmatic sclerotization highly
sclerotized, forming a subcircular crescent, each with 3 stigmatic setae, each seta 6–18
m long and bluntly
spinose or conical, subequal in length, often broken off.
Vent er. Derm membranous. Ventral setae slender, straight or slightly bent, each 10–19
m long, those on
abdominal segments usually longer, shortest around body margin. Ventral microducts with a long inner
ductule, duct rim about 2
m wide, scattered evenly on venter. Tubular ducts absent. Clypeolabral shield 216–
243
m wide. Multilocular disc-pores each 6–8 m wide with 5–7 loculi; scarce, restricted to a small group on
each side of vulva. Spiracular disc-pores each 4–5
m wide, with 5 loculi, present around spiracles and
extending to near opening of stigmatic cleft. Antennae 1 segmented, reduced to a small round sclerotized
plate; total width of plate 27–50
m. Legs represented by a remnant claw, 1 pair of digitules, several pores and
numerous small setae. Spiracles located near margin; anterior peritremes each 96–119
m wide, posterior
peritremes each 121–141
m wide.
Diagnosis. The adult female of C. reticulolaminae can be diagnosed by the following features: (i)
presence of numerous subcircular sclerotic pores arranged in a reticulate pattern, (ii) dorsal setae sharply to
bluntly spinose, with a swollen or spatulate apex, (iii) dorsal microducts heavily sclerotized, with a deep outer
ductule broadened near duct opening, (iv) preopercular pores absent, (v) presence of about 4 setae on surface
of each anal plate, (vi) marginal setae not differentiated from dorsal setae, (vii) stigmatic sclerotization each
with 3 stigmatic setae, (viii) antennae 1 segmented, reduced to a small round sclerotized plate (ix) ventral
microducts with a swollen inner ductule, (x) multilocular disc-pores restricted to a small group on each side of
vulva, (xi) legs vestigial, setose, and (xii) ventral tubular ducts absent.
Remarks. C. reticulolaminae was originally collected in Panama, but it also occurs in Costa Rica and
Mexico, suggesting a much wider distribution.
First-instar nymph (Fig. 19)
Unmounted material. Not available for study.
Mounted material. Elongate oval, 631–712
m long, 415–852 m wide (n=24).
Zootaxa 2709 © 2010 Magnolia Press · 49
REVISION OF CRYPTOSTIGMA FERRIS
Dorsum. Dorsal derm membranous. Dorsal setae each 18–22 m long; about 5 pairs present in 2 mid-
dorsal longitudinal rows extending from head region towards area dorsad to between meso- and metathorax.
Trilocular pore present on each side of head near margin close to eye. Dorsal microducts each about 2
m
wide, appearing unilocular, present in about 10 longitudinal rows. Simple pores each 3–4
m wide, present
submedially and submarginally in irregular rows. Anal plates together quadrate, dorsal surface of a shingled
texture, each plate 100–113
m long, 47–64 m wide; dorsal surface with 1 seta present on mid-area of plate
near margin, plus 4 apical setae; ventral surface with 1 fringe seta. Anal ring as in generic diagnosis. Eyespots
present just above level of antennal scape.
Margin. Outline smooth. Marginal setae each sharply spinose and 20–27
m long, totalling 70–72, with
10 anteriorly between eyes and, on each side, 8 between each eye and anterior stigmatic setae, 9 or 10 between
anterior and posterior stigmatic setae, and 12–14 between posterior stigmatic setae and body apex. Stigmatic
setae each bluntly spinose and 9–15
m long; with 3 in each anterior stigmatic cleft and 4 or 5 in each
posterior cleft.
Vent er. Ventral derm membranous. Antennae 5 segmented, total length 216–270
m. Pairs of submedian
abdominal setae present on last 3 abdominal segments. Interantennal setae 1 pair. Submarginal setae slender,
with 7 pairs on abdomen, 1 pair between each anterior and posterior stigmatic areas, and 1 pair on head
region. Ventral microducts each about 2
m wide, with 2 on each side between each antennal scape and
margin, 1 on each side anterior to anterior stigmatic areas; 5 between each anterior and posterior stigmatic
setae, and 7 between each posterior stigmatic setae and body apex. Spiracular disc-pores each 3–4
m wide
with 3–6 (mostly 5) loculi; with 4 or 5 pores in each anterior stigmatic furrow and 5–7 in each posterior
furrow. Spiracular peritremes each 11–13
m wide. Clypeolabral shield 117–149 m wide. Legs well
developed; trochanter with a very long seta, trochanter + femur 119–134
m long, tibia + tarsus 128–134 m
long; microctenidia present on apex of tibia. Tarsal digitules similar, slender. Claw without a denticle; claw
digitules slender, knobbed.
Diagnosis. The first-instar nymph of C. reticulolaminae can be diagnosed by the following combination
of features: (i) dorsal setae present in 2 longitudinal rows of 5 pairs, (ii) each anterior stigmatic cleft with 3
stigmatic setae; each posterior cleft with 4 or 5, (iii) with 9 or 10 marginal setae between anterior and posterior
stigmatic setae, (iv) antennae 5 segmented, (v) with pairs of submedian abdominal setae on last 3 abdominal
segments, (vi) anterior stigmatic furrows each with 4 or 5 pores, posterior furrows each with 5–7 pores, (vii)
ventral microducts with a long inner ductule, and (viii) dorsal surface of anal plates of a shingled texture. C.
reticulolaminae is the only species in the genus with first-instar nymphs that have ventral microducts with a
very long inner ductule.
Remarks. Despite the resemblances of adult female C. reticulolaminae to those of Myzolecanium, the
first-instar nymphs of C. reticulolaminae are typical of Cryptostigma, with 5-segmented antennae (5- or 6-
segmented antennae in Cryptostigma, and always 6-segmented antennae in Myzolecanium); absence of setae
near each coxa (a seta always present next to each coxa in Myzolecanium); presence of 1 or 2 membranous
folds just anterior to anal plates and with folds showing some sign of sclerotization (no membranous folds just
anterior to anal plates in Myzolecanium); and with 3 pairs of ventral submedian setae posteriorly on abdomen
(usually 3 pairs in Cryptostigma but always 6 pairs in Myzolecanium).
Host plants. Boraginaceae: Cordia alliodora; Polygonaceae: Triplaris americana.
Associated Hymenoptera. Formicidae: Pseudomyrmecinae: Pseudomyrmex triplaris.
Males. Unknown.
Distribution. Neotropical Region: Costa Rica, Mexico, Panama.
Cryptostigma rhizophilum Kondo, sp. nov.
(Figs 20 & 21)
Material examined. Holotype. Adult
. PANAMA (AS CANAL ZONE): Hawaii, 8.i.1957, coll. M.
Bolosan, (Hon. 38496), ex Persisteria elata, 1 (2: holotype clearly indicated on label) (USNM). Paratypes.
KONDO
50 · Zootaxa 2709 © 2010 Magnolia Press
COLOMBIA: Salado, -.v.1943, coll. S.B. Losada, (Losada # 5-158), ex pineapple roots, 1 (1) (USNM).
COSTA RICA: (Quarantine at San Francisco, California), 29.iii.1939, coll. not given, ex Zingiber officinale,
1 (1) (USNM); Intercepted at San Francisco, 29.iii.1939, coll. R.D. Clemens, (San Francisco 16254), ex abaca
(Musa textiles), 2 (4), (USNM); Bataan, 8.v.1952, coll. C.H. Batchelder, ex abaca, 2 (2) (USNM); Bataan,
22.x.1952, coll. C.H. Batchelder, (#54), ex Abaca, 2 (6) (USNM); Bataan, 3.x.1952, coll. C.H. Batchelder,
(#55), ex abaca, 2 (7) (USNM); Bataan, 26.vi.1952, coll. C.H. Batchelder, (#42), ex abaca, 2 (21: 3 adult
+
18 first-instar nymphs) (USNM); Bataan, 26.vi.1952, coll. C.H. Batchelder, (#41), ex Anthurium sp., 2 (4),
(USNM). ECUADOR: Quininde, 70 km NW Santo Domingo, date not given, coll. W.M. Allen, ex oil palm
roots, 1 (2) (BME); Quininde, 70 km NW of Santo Domingo, 5.ix.1975, coll. W.M. Allen, ex oil palm roots, 2
(16 first-instar nymphs) (BME); (intercepted) at Miami (23804), 7.iv.1980, coll. R. Faircloth, ex African palm
(Elaeis guineensis), 1 (1) (USNM); Santo Domingo, Quininde Hwy., 28 km, Hacienda la Merced of Fidel
Egas, 1.xii.1960, coll. G. Merino, ex Elaeis guineensis roots, 1 (5) (USNM); Estación Experimental Santo
Domingo, 6.vi.1983, coll. H. Vera, ex Elaeis guineensis roots, 3 (3) (USNM); Estación Experimental Santo
Domingo, 12.vi.1983, coll. H. Vera, ex Arecaceae roots, 1 (1) (USNM); Pichilinque, 1.x.1944, coll. E.J.
Hambleton, (#51), ex banana roots, 1 (1) (USNM); Ecuador, Pichilinque, 1.x.1944, coll. E.J. Hambleton,
(#55), ex orquid root, 1 (2), (USNM); Pichilinque, 1.x.1944, coll. E.J. Hambleton (#62), ex cacao roots, 1 (2
first-instar nymphs) (USNM); Santo Domingo de los Colorados, 3.x.1958, coll. G. Merino, Letter Oct. 17 (#
X-L-31), ex Peristeria elata, 1 (4) (USNM); place not given, 25.x.1995, coll. K. Ortega/M. Kaae, host not
given, LB 004673CA, 1 (1) (USNM); Santo Domingo de los Colorados, -.vi.1975, coll. Desmier, No. 6245-
(2–5, 7, 8), ex oil palm roots, 6 (11: 3 adult
+ 1 second-instar + 7 first-instar nymphs) (MNHN).
PANAMA (AS CANAL ZONE): 24.vi.1977, coll. R. Lyle, (Miami 16370), ex pineapple roots, 2 (5)
(USNM).
Adult female (Fig. 20)
Unmounted material. Live specimens were not available.
Mounted material. Body outline oval to elongate oval, 1.9–4.3 mm long, 1.6–4.0 mm wide (n=58).
Dorsum. Derm membranous, with numerous circular sclerotic pores scattered evenly on dorsum; each
pore 11–36
m wide. Dorsal setae sharply or bluntly spinose, occasionally with a swollen apex, each 8.5–13.0
m long, present evenly on dorsum. Simple pores circular to subcircular, each about 3–4 m wide, present in
subcircular clusters of 16–50 pores just laterad to stigmatic cleft. Dorsal microducts bilocular, each about 3–4
m wide; numerous, evenly distributed on dorsum. Preopercular pores absent. clerotic crescent present
around anal plates. Anal plates together quadrate, with rounded angles, each plate 146–173
m long, 76–86
m wide, anterolateral margin 110–150 µm long, posterolateral margin 85–125 µm long; with about 5 setae
on dorsal surface, plus about 1 ventral subapical setae on each plate; plates located about 1/5 of body length
from posterior margin. Anal ring with 10 setae. Eyes absent.
Margin. Margin smooth. Marginal setae sharply spinose, each 9–15
m long, arranged in a single row,
with about 14–30 setae between anterior and posterior stigmatic areas. Stigmatic clefts very deep; stigmatic
sclerotization incorporating many spiracular disc-pores. Stigmatic setae (not illustrated) each 4–11
m long
and bluntly spinose or conical, with 3 per cleft, all subequal in length, but setae often broken off or absent.
Vent er. Derm membranous. Ventral setae slender, straight or slightly bent, each 13–23
m long, those on
abdominal segments usually longest. Ventral microducts each about 3
m wide, scattered evenly on venter.
Tubular ducts absent. Clypeolabral shield 243–291
m wide. Multilocular disc-pores each 5–7 m wide with
5–10 (mostly 7) loculi, present on perivulvar region. Spiracular disc-pores each 4–6
m wide with 3–7
(mostly 5) loculi, present from area of spiracles to body margin. Antennae very small, 1 segmented,
represented by a flattened segment bearing several setae, length 17–21
m. Interantennal setae 1 pair. Legs
greatly reduced, with segments fused and forming a sclerotized disc, total length 21–32
m, each leg bearing
a claw, a pair of hair-like digitules and numerous setae. Claw without a denticle. Spiracles larger than legs,
located near margin; each anterior peritremes 70–83
m wide, each posterior peritremes 72–83 m wide.
Zootaxa 2709 © 2010 Magnolia Press · 51
REVISION OF CRYPTOSTIGMA FERRIS
FIGURE 20. Cryptostigma rhizophilum Kondo, adult female.
KONDO
52 · Zootaxa 2709 © 2010 Magnolia Press
FIGURE 21. Cryptostigma rhizophilum Kondo, first-instar nymph.
Zootaxa 2709 © 2010 Magnolia Press · 53
REVISION OF CRYPTOSTIGMA FERRIS
Diagnosis. The adult female of C. rhizophilum can be diagnosed by the following features: (i) sclerotic
pores present, (ii) dorsal setae sharply or bluntly spinose, occasionally with a swollen apex, (iii) preopercular
pores absent, (iv) each anal plate with about 5 setae on dorsal surface, (v) with about 14–29 marginal setae
between anterior and posterior stigmatic areas, (vi) antennae 1 segmented, reduced to a small round plate
bearing numerous setae, (vii) multilocular disc-pores restricted to perivulvar region, (viii) legs greatly
reduced, with segments fused and forming a sclerotized disc bearing a claw and numerous setae, and (ix)
ventral tubular ducts absent. C. rhizophilum is the only known species in the genus with subcircular clusters of
simple pores present laterad to each stigmatic cleft.
First-instar nymph (Fig. 21)
Unmounted material. Not available for study.
Mounted material. Nymphs elongate oval, 706–792
m long, 501–555 m wide (n=36).
Dorsum. Dorsal derm membranous. Dorsal setae each 7–16
m long, totalling about 5 pairs, present in 2
mid-dorsal longitudinal rows, extending from head region towards area dorsad to between meso- and
metathorax. Trilocular pore present on each side of head near margin close to eye. Dorsal microducts each
about 2
m wide, appearing bilocular, present in about 8 longitudinal rows. Simple pores each 2–4 m wide,
present submedially and submarginally. Anal plates together quadrate, each plate 216–237
m long, 108–124
m wide; dorsal surface with 1 seta present on mid-area of plate near inner margin, plus 3 apical setae; ventral
surface with 1 fringe seta. Anal ring as in generic diagnosis. Eyespots present on margin just above level of
antennal scape.
Margin. Outline smooth. Marginal setae sharply spinose, each 15–23
m long, totalling 75–80, with 10–
13 anteriorly between eyes and, on each side, 7–11 between each eye and anterior stigmatic setae, 8–13
between anterior and posterior stigmatic setae, and 14–16 between each posterior stigmatic setae and body
apex. Stigmatic setae each 11–18
m long and bluntly spinose; with 3 setae in each anterior stigmatic cleft and
3 or 4 in each posterior cleft.
Venter. Ventral derm membranous. Antennae 5 segmented, total length 216–248
m. Mid-ventral setae
slender, with a pair in posterior 3 abdominal segments. Interantennal setae 1 pair. Submarginal setae of 2
types: inner submarginal setae slender, present in 1 or 2 rows; outermost submarginal setae sharply spinose,
present in 1 row on abdomen between posterior stigmatic areas and body apex, with 4 sharply spinose setae
between each anterior and posterior stigmatic areas, and 1 pair of slender setae on head region. Ventral
microducts each about 2
m wide, with 4 on each side between each antennal scape and anterior stigmatic
area, 4 between each anterior and posterior stigmatic setae, and 5 or 6 between each posterior stigmatic area
and anal cleft. Spiracular disc-pores each 3.5–4.0
m wide with 3–8 (mostly 5) loculi; both anterior and
posterior stigmatic furrows each with 5–8 pores. Spiracular peritremes each 12–15
m wide. Clypeolabral
shield 145–160
m wide. Legs well developed, trochanter with a very long setae, trochanter + femur 280–313
m long, tibia + tarsus 345–367 m long; microctenidia present on apex of tibia. Prothoracic tarsal digitules
dissimilar, 1 knobbed and 1 spiniform, meso- and metathoracic tarsal digitules similar, knobbed. Claw without
a denticle; claw digitules slender, knobbed.
Variation. In the first-instar nymphs of C. rhizophilum, the submarginal setae between the posterior
spiracle and anal cleft are present in either 2 or three parallel rows, the outermost row being composed of
sharply spinose setae.
Diagnosis. The first-instar nymph of C. rhizophilum can be diagnosed by the following combination of
features: (i) dorsal setae present in 2 longitudinal rows of 5 pairs, (ii) each anterior stigmatic cleft with 3
stigmatic setae; posterior stigmatic cleft with 3 or 4 setae, (iii) 8–13 marginal setae present between anterior
and posterior stigmatic setae, (iv) antennae 5 segmented, (v) with pairs of submedian abdominal setae on last
3 abdominal segments, (vi) each anterior and posterior stigmatic furrow with 5–8 pores, and (vii) 1 microduct
present mesad to each inner submarginal setae on abdomen. C. rhizophilum is the only species in the genus
with first-instar nymphs that have sharply spinose outer submarginal setae.
Remarks. C. rhizophilum is unusual among species of Cryptostigma in its preference for the roots of its
host. Two other species of Cryptostigma have been recorded from roots: C. urichi, a commonly ant-tended
scale insect, normally found inside hollow branches and stems, has been collected on the large roots of
KONDO
54 · Zootaxa 2709 © 2010 Magnolia Press
Erythrina sp. in Surinam, although these were probably the exposed larger roots of the tree and not the
underground roots; and C. silveirai appears to be restricted to the underground roots of grape vines and is
known to be an important pest of vines in Brazil (see remarks under C. silveirai). C. rhizophilum has the
potential to become an agricultural pest since its hosts includes oil palm, banana, Manila hemp, orchids and
pineapple.
Host plants. Araceae: Anthurium sp. Arecaceae: Elaeis guineensis. Bromeliaceae: pineapple (Ananas
sp.). Musaceae: Musa sapientum, Musa textilis. Orchidaceae: Peristeria elata. Sterculiaceae: Theobroma sp.
Zingiberaceae: Zingiber officinale.
Associated Hymenoptera. None reported.
Etymology. The specific epithet “rhizophilum” is composed of the Greek words “rhiza” meaning “root
and “philum” which is a neutral adjective meaning loving.
Males. Known from second-instar male nymphs.
Distribution. Neotropical Region: Colombia, Costa Rica, Ecuador, Panama.
Cryptostigma saundersi Laing
(Fig. 22)
Cryptostigma saundersi Laing: Laing 1925: 59; Ben-Dov 1993: 99; Qin & Gullan 1989: 225.
Material examined. Lectotype, here designated. Cryptostigma saundersi Laing, adult . BRAZIL: Rio de
Janeiro, ex trunk of a tree protected by ants, 9.viii.1923, coll. L.G. Saunders (# 4), 1 (1) (BMNH).
Paralectotype. Same data as lectotype, except that no date given, 1 slide with 1 specimen with dorsum and
venter under separate cover slips (BMNH).
Adult female (Fig. 22)
Unmounted material. Alcohol preserved specimens with a pale yellowish brown longitudinal stripe and
several intertwining stripes on dorsum; also with semicircular stripes on declivity anterior to anal lobes;
remainder of dorsum of a darker brown; a small darker brown to almost black area around anal plates. Ventral
surface dark brown with a paler marginal area. Length (unmounted) approximately 7 mm, breadth from 5–6
mm (Laing, 1925).
Mounted material. Body outline oval, constricted at stigmatic areas, 6.0–7.0 mm long, 6.5–7.0 mm wide
(n=2).
Dorsum. Derm membranous, with numerous (68–92) subcircular compound orbicular pores, scattered on
dorsum, size variable, those closest to body margin smallest, abundant on body margin and submargin but
absent from mid-dorsum; each orbicular pore associated with 4–15 bilocular microducts, and about 6–24
slender setae around pore margins; each seta 13–19
m long. Dorsal setae absent, except for those on margins
of orbicular pores. Simple pores of 2 sizes: larger pores each about 8
m wide; smaller pores each about 4.5
m wide, scattered throughout dorsum. Dorsal microducts of 2 types: small microducts each about 2 m wide
and unilocular; larger microducts each about 6.0
m wide and appearing bilocular, both types scattered on
dorsum. Preopercular pores absent. Narrow sclerotic crescen