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63
Proceedings of the XI International Symposium on Scale Insect Studies
The morphology of lac insects (Hemiptera: Coccoidea: Kerriidae)
P J Gullan & T Kondo
Department of Entomology, University of California, One Shields Ave., Davis, CA 95616-8584,
U.S.A.
Abstract: Lac insects constitute a family, the Kerriidae, of morphologically peculiar scale insects that
produce resinous secretions that mostly form a hard scale cover or test. Identification to species level
is based on membranous, globose adult females that are difficult to slide-mount and have a complex
morphology that is unfamiliar even to most coccidologists. Characteristics important to species
recognition are poorly understood in most genera, and the species diversity of lac insects probably
has been underestimated. Here we present an overview of the cuticular morphology of the Kerriidae,
with emphasis on features most important for species identification, such as the anal tubercle,
brachial plate, dorsal spine and marginal and ventral duct clusters. We provide a detailed illustration
of the young adult female of Kerria lacca (Kerr), the lac insect of commerce.
Key words: Kerriidae, lac insects, morphology, Kerria lacca.
Introduction
Approximately 90 species of lac insects (Hemiptera: Coccoidea: Kerriidae) have been
described worldwide in nine genera (Ben-Dov, 2006), based on a classification that was established
by the monographic work of Joseph C. Chamberlin (1923, 1925). The most widely known lac insect
species, Kerria lacca (Kerr), is renowned for the commercial use of its secretion, or shellac, as a
natural polymer (Varshney, 1970; Yao et al., 1989; Ben-Dov, 2006). Some other lac insect species
are plant pests that can cause branch dieback or even plant death (e.g., Campbell et al., 1994;
Pemberton, 2003). Although taxonomic studies in China (e.g., Wang, 1986; Zhang, 1992, 1993),
India (e.g., Varshney, 1977, 1984) and South Africa (Munting, 1965, 1966) have furthered our
knowledge of kerriid diversity, and recent research by Ireneo L. Lit, Jr (Lit & Gullan, 2001; Lit,
2002a,b) has increased our understanding of basic lac insect morphology, accurate identification to
species level remains challenging for several reasons. First there is the technical difficulty of
preparing good microscope slide-mounts that are essential for seeing some of the diagnostic
cuticular features. Adult females have globular, membranous bodies and, unless correctly flattened
dorsoventrally, it may be impossible to distinguish or count structures such as the number of
microducts in marginal and ventral duct clusters. Second, there is an idiosyncratic nomenclature
associated with the peculiar and complex morphology of adult female lac insects, and this can be a
deterrent to using keys or taxonomic papers. Third, we have a very poor understanding of what
constitutes a lac insect species. There are no molecular phylogenetic or population genetic studies
of lac insects, not even for K. lacca and its relatives, and our recent taxonomic study of
Paratachardina Balachowsky (Kondo & Gullan, 2007) has shown that the adult females of closely-
related species can be extremely similar in their cuticular morphology. It is likely that there are
many undescribed species of kerriids.
Here we review the morphological features of the adult female lac insects, provide the first
detailed illustration of the young adult female of K. lacca (the type species of the type genus Kerria
Targioni Tozzetti), and discuss cuticular structures of taxonomic importance in the identification of
kerriid species.
Material and Methods
Young adult females of K. lacca were slide-mounted from dry material deposited at the
BME (Bohart Museum of Entomology, University of California, Davis). The collection
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Proceedings of the XI International Symposium on Scale Insect Studies
information for these specimens is as follows: "Tachardia lacca grown in Ceylon from imported
stock on Albizzia sp., Peradeniya, Ceylon, # 8940, E.E. Green", 11 slides, 11 specimens, slide-
mounted by T. Kondo and P.J. Gullan, 2007. The illustration of K. lacca is a generalization of
several specimens and was prepared with the assistance of a camera lucida attached to an Olympus
BX40 compound microscope. The terms used to describe lac insects follow those of Chamberlin
(1923), Gill (1993) and Kondo & Gullan (2005, 2007). Some important structures are illustrated in
Figure 1, and others are shown in Figure 2.
Results and discussion
Lac insect females have three instars, i.e., first (crawler stage), second, and adult. Males, if present,
have five instars, i.e., first, second, prepupal, pupal and adult. Although the current classification of
lac insect genera and species is based almost entirely on the cuticular morphology of the adult
females, in the future other instars, especially first-instar nymphs, may become useful for
identification. For a generalized drawing of a lac insect first-instar nymph, see Miller (1991); for a
description of the first-instar nymph of K. lacca see Misra (1931); for the first- and second-instar
nymphs of a Paratachardina species, see Kondo & Gullan (2007).
Lac insects are a morphologically distinctive group of scale insects characterized by adult
females with the following features: (i) body generally covered in a resinous test; (ii) dorsal spine
present (absent in only one species); (ii) brachial plates present; (iii) anal tubercle present and
usually composed of a sclerotized supra-anal plate and usually a less sclerotized pre-anal plate; (iv)
anal ring with 10 setae; (v) antennae with 1–7 segments; (vi) legs reduced or absent; (vii) anterior
spiracles located on body margin or on dorsum, and much larger than posterior spiracles; (viii)
canella (anterior spiracular furrow lined with pores) present or absent; (ix) ventral setae present,
usually short and scarce; (x) marginal duct clusters present; (xi) ventral duct clusters usually
present; (xii) spermatoid ducts present on both surfaces of body: on venter restricted generally to
marginal and submarginal areas and most abundant within each marginal duct cluster, and on
dorsum more-or-less widely distributed throughout, but usually with areas devoid of ducts, e.g.,
around dorsal spine, anal tubercle and brachial plates. Within a species, the body shape of the adult
female can vary considerably (see Takahashi (1941) for illustration of this variation in K. lacca),
depending on age and growth conditions.
The adult female resembles the second-instar female, but differs as follows (character states of
second instar in parentheses if appropriate): (i) larger body size; (ii) with many more ducts and
pores; (iii) larger brachial plates; (iv) well developed brachia (absent in the second instar); (v)
almost always with a dorsal spine (absent or represented by a membranous lobe); and (vi)
perivulvar pores present in some taxa (always absent). Besides their smaller size, first-instar
nymphs are readily differentiated from subsequent instars by: (i) presence of one pair of very long
setae on the anal tubercle, each about half length of body; (ii) well developed antennae with 6
segments; (iii) cursorial legs; and (iv) lack of brachial plates and brachia (Misra, 1931; Miller,
1991; Kondo & Gullan, 2007).
Adult female: the resinous test
Typically the lac test is composed of a hard resin, although the test is described as sticky in
Austrotachardiella sexcordata Matile-Ferrero (Matile-Ferrero & Couturier, 1993). The tests of
many lac insect taxa dissolve in alcohol, whereas tests of other taxa, such as Paratachardina
species, do not dissolve (Varshney, 1984). The tendency for some tests to dissolve more
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Proceedings of the XI International Symposium on Scale Insect Studies
readily than others may have lead to the idea that some tests are gum-like and pliable, but there is
no comparative study of lac insect tests, and the chemistry of very few tests have been studied. The
test greatly varies in shape and color, and often the tests of crowded individuals fuse to form a
single mass. The test of a single individual may be unlobed and rounded, or with 3–6 marginal
lobes. The surface of the test may be smooth or with a series of longitudinal ridges, and often
various bumps or elevations may be present. According to Green (1922), the shape of the insect
body can be reflected in the shape of its test.
Lac tests vary from dull brown or dark, to various tones of red or orange, although species with
greenish or yellowish colors also exist. Each test is usually of a single colour, but some species may
have characteristic color patterns, e.g., Paratachardina decorella (Maskell).
Adult female: cuticular structure
Dorsum. The derm is always membranous, and without visible segmentation. Dorsal setae
(dset) generally are short and scarce, but usually are present around the body margin and often there
are a few around the dorsal spine and dorsal tubercle. One of the most distinctive features of lac
insects is a pair of brachial plates (brpl), each often elevated on a membranous to sclerotized,
tubular protrusion called a brachium (br) [see Lit (2002a) for a discussion of other terms used for
this structure]. Characteristics of the brachium and brachial plate have been used extensively for
distinguishing species within genera (e.g., Chamberlin, 1923; Varshney, 1977; Kondo & Gullan,
2007) and for separating some genera (Chamberlin, 1923; Zhang, 1992; Kondo & Gullan, 2005).
Each brachial plate is heavily sclerotized, and depending on species, can vary in shape from round,
triangular, quadrate to irregular in outline, and may be flat or have a shallow or deep crater, or an
elevated protrusion. Each plate bears brachial pores, brachial plate setae and pseudospines, the
number of which can be informative for separating some species of Paratachardina (Kondo &
Gullan, 2007), although variation within a species can confound taxonomic utility. Brachial pores
usually have 5 loculi and are similar in structure to spiracular pores and canellar pores. A
pseudospine is a specialized type of multilocular pore that resembles a seta or spine when viewed
from the side, but a pore with 5 (rarely 3, 4 or 6 loculi) when viewed from above (Chamberlin,
1923; Lit 2002a,b). In the first-instar nymphs, the homologous structures are hollow spines located
in the anterior spiracular cleft and in the pseudocerarii (Miller, 1991). The brachia either can be
absent, very short (sessile) or very long. Typically, the base of each brachium is associated closely
with one of the large anterior spiracles; secretions from the brachial plate are believed to keep the
spiracular openings of the test from being sealed over by the resin of the test (Chamberlin, 1923).
The dorsum is characterised by the presence of another unique structure, the dorsal spine (dspi),
which is absent only in Tachardina albida Cockerell (Chamberlin, 1923). This conical, hollow
spine is located anterior to the anal tubercle, and its apex probably always has an opening that
connects to an internal duct, called the dorsal spine duct. The duct length and structure varies
greatly among species, but ducts usually are divided into two types: (i) the dendritic type is most
common and is quite branched throughout its entire length, as in Austrotachardiella colombiana
Kondo & Gullan (Fig. 1A) and K. lacca (Fig. 2); and (ii) the non-dendritic (= fibrous) type
is only known in Austrotachardia Chamberlin and is unbranched throughout its length,
either ending on a swollen or a branched apex as in A. melaleucae (Maskell) (Fig. 1B)
(Lit, 2001a). The dorsal spine is often elevated on a
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Proceedings of the XI International Symposium on Scale Insect Studies
membranous tube, called the pedicel, that can be very short or absent, or very long and slender as in
Albotachardina species. The function of the dorsal spine and its duct is unknown (see Lit 2002a for
discussion of previous hypotheses), but any hypothesis must be able to account for lack of
development of the dorsal spine in first- and second-instar nymphs (the latter are collected rarely
and probably short-lived).
Figure 1. A, dorsal spine and dendritic spine duct of the adult female of Austrotachardiella
colombiana Kondo & Gullan; B, dorsal spine and non-dendritic spine duct of the adult female of
Austrotachardia melaleucae (Maskell); C, duplex type of marginal duct cluster as in Tachardiella
species; D, triplex type of marginal duct cluster as in Austrotachardiella species; E, marginal duct
cluster with two associated auxiliary clusters as in Afrotachardina brachysetosa (Chamberlin); F,
typical macroduct of Austrotachardiella species.
The anal tubercle (at) is another characteristic feature of lac insects, although in some members of
the Coccidae that have a thick waxy covering, such as Ceroplastes Gray, the area surrounding the
anal plates can be highly sclerotized and of a tubular shape somewhat similar to that seen in the
Kerriidae. In lac insects, the anal tubercle is the posterior sclerotized prolongation of the body that
bears the pre-anal plate (papl) (in some groups), the supra-anal plate (spanpl), the anal fringe and
the anal ring (Fig. 2). The anal fringe (af) is a series of ligulate lobes at the apex of the anal tubercle
encircling the anal ring. The anal ring of kerriids may be divided into 4, 6 or more distinct sections
or it can be entire (unsegmented) or show partial segmentation, and it always has 10 setae and
usually an irregular row of pores. Some species of Paratachardina and Tachardina Cockerell have
pygidial apodemes, which are variably sclerotized internal rods or processes that extend from the
base of the supra-anal plate posteroventrally. Chamberlin (1923:158) incorrectly stated that these
apodemes extended to "caudad of the dorsal spine". They probably function as rigid
attachments for muscles that move the anal tubercle or its components.
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Proceedings of the XI International Symposium on Scale Insect Studies
Figure 2. Kerria lacca (Kerr), young adult female (see text for abbreviations).
Microducts are either more or less scattered evenly throughout dorsum, or present only around
the margins and submargins, and often there are fewer or absent around the anal tubercle, anterior
spiracles, brachial plates and dorsal spine (Fig. 2). Microducts of the Kerriidae resemble the typical
coccid ventral microduct, with a subcircular duct opening and short terminal gland. Spermatoid
ducts (spd) (= spermatozooid ducts), which are unique to lac insects, are long thread-like ducts with
a greatly swollen area at the inner end distal from the duct opening (Fig. 2). These ducts may occur
singly on the derm or, in some genera, associated with the rim of microducts, and may contribute to
the resin of the lac test. For a discussion of the naming of this duct, see Kondo & Gullan (2007).
The Margin. Unlike most other scale insects, the margins of lac insects are poorly defined and,
in many species, it is hard to differentiate the dorsum from the venter after the adult female
becomes mature and fully expanded (e.g. K. lacca). However, it is possible to locate the marginal
areas by the presence of short setae or by the distribution of marginal duct clusters (mdc). The
presence of mdcs is another distinctive characteristic of lac insects. Marginal duct clusters are
subcircular to irregular clusters of microducts and / or macroducts
and / or spermatoid ducts present around the body margin, usually on the venter, but
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Proceedings of the XI International Symposium on Scale Insect Studies
sometimes apparently on the dorsum (e.g. K. lacca). In Afrotachardina Chamberlin, mdcs have two
closely associated auxiliary clusters of microducts (Fig. 1E). The typical microducts of most lac
insects can be characterised arbitrarily as small-sized, medium-sized (msm) or large-sized (lsm) (Fig.
2). The structure of the different size classes of microducts generally are the same, however, in the
New World genera Austrotachardiella Chamberlin and Tachardiella Cockerell, there are large-
sized ducts, called macroducts (Kondo & Gullan 2005, 2007), of a different structure to the smaller
microducts. Macroducts have a cylindrical shape with a long and slender tail that splits near its end
(Fig. 1F). The presence or absence, sizes, and combination of microducts, macroducts and
spermatoid ducts in the mdcs can be useful features for identification at both the genus and species
level. For example, the number of macroducts per marginal duct cluster is useful in separating
species of Austrotachardiella and Tachardiella (Chamberlin 1923; Kondo & Gullan 2005).
Chamberlin (1923) describes a series of types of marginal duct clusters. His simplex mdc consists
of ducts of one size and structure, as in K. lacca (Fig. 2), his duplex mdc contains ducts of two
types or sizes but with many variants of duct structure, whereas the triplex type of mdc has three
kinds of ducts. In Tachardiella, duplex mdcs are composed of macroducts and spermatoid ducts
only (Fig. 1C), whereas Austrotachardiella species have triplex mdcs composed of macroducts,
spermatoid ducts and an outer narrow band of microducts (Fig. 1D). Chamberlin's classification
into simplex, duplex and triplex mdcs fails to consider the homologies of duct types. For example,
the name duplex mdc would be applied to an mdc with large central and small outer microducts, as
well as to a mdc composed of macroducts and spermatoid ducts. Furthermore, Chamberlin (1923)
confusingly refers to the central larger ducts of any type of mdc as "nuclear ducts", and to all other
ducts in a cluster as "duplex ducts" (meaning secondary ducts), regardless of structure. The nature
and function(s) of secretions from mdcs are unknown, but the secretions must contribute to the lac
test.
The Venter. The derm is membranous, with segmentation sometimes visible on medial areas
of the thorax and abdomen. Microtrichia are often present in segmental transverse rows in these
areas, and usually a few setae (vset) are present on each of these segments. The antennae (ant) are
always present and have 1–7 segments, although the segmentation is often poorly defined. Antennal
position may vary from slightly anterolateral to the mouthparts, to near the body margin lateral to
the mouthparts (as in Paratachardina), or displaced onto the dorsal surface of the head. The
mouthparts of lac insects have a labium of 1–2 segments and generally 5–6 pairs of labial setae
(Koteja 1974). In some taxa, there are one or two pairs of membranous, often slightly sclerotized
lobes surrounding the mouthparts. The lobes posterior to the mouthparts are called the post-oral
lobes (pol) (Chamberlin 1923) and more anterior lobes, if present, are called the pre-oral lobes (prl)
(Kondo & Gullan 2007). In the Kerriidae, as in all neococcoid scale insects, there are two pairs of
thoracic spiracles and no abdominal spiracles. The anterior spiracles (antsp) of the adult female may
be located either marginally or on the dorsum and always are larger than the posterior spiracles
(postsp), which are present typically on the venter (Fig. 2). The multilocular pores present
surrounding the spiracular peritreme are called spiracular pores (spp) (vignette Fig. 2). These pores
mostly have 5 loculi and are similar in structure to canellar pores and brachial pores. The term
canella (plural canellae) was coined by MacGillivray (1921) for the spiracular furrow
of scale insects, but was adopted later (Chamberlin, 1923) for the linear group of
multilocular pores [canellar pores (canp)] between the anterior spiracles and
mouthparts, or between the anterior and posterior spiracles as in some
Tachardiella species. Recently Kondo & Gullan (2007) expanded the definition to
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Proceedings of the XI International Symposium on Scale Insect Studies
include the multilocular pores present between the anterior spiracles and brachial plates in
Paratachardina.
Multilocular pores found around the vulva are called perivulvar (= perivaginal) pores (pvp),
and are found generally within perivulvar (= perivaginal) pore clusters (pvpc), although they may
occur singly (as in Tachardina species). Perivulvar pore clusters can be absent entirely in some
genera (e.g. Austrotachardia and Paratachardina), or range from 4 in Tachardiella and
Austrotachardiella to 18–50 in Kerria. Microducts usually vary in size within a species (as
described for the dorsum), but microducts scattered on the derm often are smaller than those in the
marginal and ventral duct clusters. The ventral duct clusters (vdc) are subcircular to irregular-
shaped clusters of microducts present submarginally or submedially around the ventral body, and
often lie parallel to the marginal duct clusters. The number of pairs of vdcs and the number of
microducts per cluster are useful for separating species of some genera, e.g. Paratachardina
(Kondo & Gullan 2007). Spermatoid ducts, similar to those on the dorsum, usually are present
marginally and submarginally on the derm but are most abundant within each marginal duct cluster
and often also within the ventral duct clusters, and generally are absent from medial areas of the
venter.
Acknowledgements
Thanks to Peter Cranston and Dug Miller for helpful comments on a draft of the manuscript. This
work was supported by the U.S. National Science Foundation (Partnership for Enhancing Expertise
in Taxonomy program, under Grant No. 0118718) and by Hatch funding from the California
Agricultural Experiment Station.
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To be cited as: Gullan, P.J. & Kondo, T. 2008. The morphology of lac insects (Hemiptera: Coccoidea: Kerriidae) 63–70 In:
Branco, M., Franco, J.C. & Hodgson, C.J. (Editors), Proceedings of the XI International Symposium on Scale Insect Studies,
Oeiras, Portugal, 24–27 September 2007. ISA Press, Lisbon, Portugal. 322 pp. *THIS COPY IS NOT AN ORIGINAL.
