Chapter 2
Morphological Adaptations of Parasitic
Arthropods
Nathan D. Burkett-Cadena
University of Florida, Florida Medical Entomology Laboratory, Vero Beach, FL, United States
There are several independent lineages of arthropods that
are parasitic on vertebrates in one or more stages of their
life cycles (Fig. 2.1). Many morphological features of these
parasitic arthropods are modified in a number of ways as
adaptations for parasitic relationships. Common adapta-
tions include those for feeding on vertebrate blood and
other body fluids, attaching to and clinging to hosts, and
dispersing to new hosts. Morphological structures that play
a particularly important role in host associations are body
shape, mouthparts, and legs. These structures are often
highly modified in parasitic arthropods. Other features that
are often modified in parasitic arthropods are the wings,
eyes, and various sensory organs.
BODY SHAPE AND WINGS
A recurring theme among ectoparasites is modification of
the general body shape to facilitate movement on the host
and to enable them to hide in tight spaces when off the host.
For parasitic lineages that exhibit long-term, close associ-
ation with host animals, this usually involves dorsoventral
or lateral flattening of the body. Dorsoventral flattening is
characteristic of bed bugs and bat bugs, lice, beaver beetles
(Leiodidae, genus Platypsyllus), parasitic dermapterans
(genus Hemimerus), louse flies and keds, and ticks. In
contrast, lateral flattening is best exemplified by fleas.
Blood-feeding arthropods that have only brief contact with
hosts, such as skin-piercing moths, mosquitoes, and biting
midges, generally do not exhibit lateral or dorsoventral
compression of the body.
Wings are indispensable features of many parasitic in-
sects, without which they would not be able to reach their
hosts. Tabanids (horse flies), for example, are strong fliers;
some species may attain air speeds of 45 km/h or greater. In
other parasitic insects, however, wings have become
secondarily reduced, or even completely lost, as in fleas,
Medical and Veterinary Entomology. https://doi.org/10.1016/B978-0-12-814043-7.00002-9
Copyright © 2019 Elsevier Inc. All rights reserved.
lice, bed bugs, and the sheep ked. Still others, such as
hippoboscids of the genus Lipoptena that parasitize deer,
may have fully functional wings as adults but shed them
after reaching a suitable host. In the latter case, the wings
break off at a specific location near their base, leaving a
wing stump.
MOUTHPARTS
Mouthparts of parasitic arthropods are typically adapted for
feeding on host body fluids, particularly blood but also
lymph, skin secretions, and tears. They also may be
adapted for feeding externally on skin, sloughed skin
scales, hair, or feathers. Those arthropods that feed directly
on host tissues generally retain the chewing-type mouth-
parts like those of cockroaches (Fig. 2.2A). Fluid-feeding
parasites, on the other hand, have mouthparts adapted for
piercing host skin to reach and feed on internal fluids. It is
among this group that the mouthparts have become the
most modified and specialized. Arthropods that use their
mouthparts to lacerate host skin and feed on blood that
pools at the bite site as a result of damage to the sur-
rounding blood vessels are called telmophages. Examples
of telmophages include black flies, biting midges, horse
flies, and deer flies. Bites from these insects typically cause
more immediate pain and discomfort at the bite site due to
puncture wounds and damage to surrounding skin and
related tissues, resulting in a greater risk of secondary in-
fections. Hematophagous arthropods with highly special-
ized piercingesucking mouthparts that pierce individual
capillaries and then feed directly on host blood are called
solenophages. Examples of solenophages are mosquitoes,
bed bugs, kissing bugs, and sucking lice. They represent
the more refined blood feeders, with highly modified,
styletiform mouthparts that typically leave little or no ev-
idence of an actual puncture of the skin at the bite site.
17
18 Medical and Veterinary Entomology
FIGURE 2.1 Arthropod phylogeny illustrating relationships of parasitic
lineages. Composite image redrawn after Regier et al. (2010), Sasaki et al.
(2013), and Lozano-Fernandez et al. (2016).
Biting midges are excellent examples of telmophages.
The labrum, mandibles, maxillae, and hypopharynx are
bladelike, pointed or serrate apically, and function in
lacerating host skin (Fig. 2.2C). The labium serves as a
sheath for protecting the other, more delicate, mouthparts.
In insects with chewing mouthparts, the labium is analo-
gous to a “lower lip” (Fig. 2.2A) and functions in grasping,
manipulating, and retaining food.
Mosquitoes are classic examples of solenophages. The
labrum, maxillae, mandibles, hypopharynx, and labium are
very elongate, forming a feeding apparatus called the pro-
boscis (Fig. 2.2B). The mosquito labium serves as a pro-
tective sheath and a guide for the fascicle, the bundle of
thinner and more delicate mouthparts that penetrate the host
skin, deliver saliva, and transport host blood. The fascicle is
composed of the labrum, maxillae, mandibles, and hypo-
pharynx, all of which are in the form of interconnecting fine
stylets. The labrum, which functions as the “upper lip” of
insects with chewing mouthparts (Fig. 2.2A), forms the
blood-feeding tube in mosquitoes. The mandibles and
maxillae, typically used for manipulating and masticating
food by arthropods with chewing mouthparts (Fig. 2.2A),
are modified in mosquitoes for piercing the host epidermis.
The hypopharynx, a tongue-like structure in insects with
chewing mouthparts (Fig. 2.2A), is also styletiform in
mosquitoes and is used to pierce host tissue. Running the
length of the hypopharynx is a channel that delivers saliva
to the apical portion of the mouthparts during feeding.
In fleas, the epipharynx (an outgrowth of the body wall
unique to fleas) and the maxillae are in the form of stylets
and are used to pierce skin (Fig. 2.2D). During blood
feeding, the tip of the epipharynx is inserted into a capil-
lary. The pair of maxillae, together with the epipharynx,
forms the feeding tube. In fleas the labium is reduced, with
only the palps visible externally. The palps help to guide
the blood-feeding stylets.
In blood-feeding hemipterans, such as bed bugs and
kissing bugs, the maxillae and mandibles are styletiform
and held within a sheathlike, segmented labium (Fig. 2.2E).
Each of the paired maxillae is curved medially and in-
terlocks with the other on their dorsal and ventral surfaces
to form the food canal. The maxillae, mandibles, and
labium form the rostrum, which is directed posteriorly and
held under the head and thorax of the insect when not in
use. The labrum is relatively unmodified and resembles the
labrum of insects with chewing mouthparts.
In sucking lice the feeding apparatus is different from
that of other parasitic insects. The labrum is highly modi-
fied to form a snoutlike structure, called the haustellum,
which surrounds the other mouthparts (Fig. 2.2F). At the tip
of the haustellum are prestomal, or haustellar, “teeth,”
which are used to anchor the mouthparts to the host. The
maxillae, hypopharynx, and labium are modified as stylets
for piercing host tissues. The hypopharynx also serves as a
salivary canal, while the maxillae form the food canal.
Among the arachnids, ticks represent a highly special-
ized group of mites that are obligate parasites of terrestrial
vertebrates. Ticks feed on host blood during each of their
developmental stages (i.e., larvae, nymphs, and adults). The
mouthparts are modified for piercing host tissue, anchoring
the tick to its host and drawing blood into the alimentary
tract. In most groups of mites, the tip of each chelicera is
pincer-like, or chelate, with a fixed digit and opposable
movable digit for grasping and manipulating food
(Fig 2.3A). In ticks, however, the chelicerae have lost the
terminal chelae and have become modified as short blade-
like structures, with serrate tips, adapted for piercing host
skin (Fig. 2.3C). The hypostome, a ventral projection of the
 Morphological Adaptations of Parasitic Arthropods Chapter | 2 19

FIGURE 2.2 Head and mouthparts of medically important insects, with cross section of mouthparts. (A) Cockroach, Periplaneta (Blattidae);
(B) mosquito, Aedes (Culicidae) with labium reflexed to show styletiform mouthparts; (C) biting midge, Culicoides (Ceratopogonidae); (D) cat flea,
Ctenocephalides felis (Pulicidae); (E) bed bug, Cimex (Cimicidae); (F) human body louse, Pediculus humanus (Pediculidae). fc, food canal. Original by
Nathan D. Burkett-Cadena.

fused palpcoxae in mites, is greatly enlarged and serves as
an attachment organ in ticks. Rearward projecting teeth on
the hypostome serve to anchor the tick securely to its host
after it has cut a hole through the skin with its chelicerae.
The palps of ticks are relatively unmodified and do not
penetrate the skin of the host. In other parasitic mites,
including members of the families Dermanyssidae and
Macronyssidae, the chelicerae may be long, slender, and
retractable and are adapted for piercing host skin. The
terminal portion of the chelicera may be pincer-like, as in
Ornithonyssus (Macronyssidae) (Fig. 2.3B), or serrate,
lacking the movable element, as in Dermanyssus (Derma-
nyssidae) (Fig. 2.3B inset).
LEGS
In both insects and arachnids, those taxa that live for
extended periods of time on their hosts often have highly
enlarged legs and/or the legs bear specialized structures to
facilitate attachment to the host and movement amidst the
host hair or feathers. Structures for grasping are often
coupled with stout, heavily sclerotized and generously
muscled appendages. This combination of characters al-
lows ectoparasites not only to obtain access to new hosts
but also to avoid being displaced or removed by host
grooming.
The insect leg typically consists of five segments
(Fig. 2.4A). The basal segment is the coxa, followed by the
trochanter, femur, tibia, and tarsus. The tarsus is divided
into subsegments, or tarsomeres, providing flexibility.
Claws and other structures, when present, are typically
borne on the apical tarsomere. In many ectoparasitic insects
the typical leg has become modified to facilitate host
attachment and dispersal to new hosts. Adult fleas exhibit
perhaps the most widely recognized modification of the
legs for getting on and off hosts. The hind legs of fleas are
particularly modified to enable them to jump remarkable
distances to reach a host or to evade removal by host
grooming (Fig. 2.4B). Modifications of the flea hind leg
include an enlarged, muscular femur (as in other jumping
insects) and an elastic protein in the integument called
resilin. Resilin, an important structural component of the
20 Medical and Veterinary Entomology
FIGURE 2.3 Gnathosoma of medically important mites, ventral views. (A) Pneumolaelaps (Laelapidae); (B) Ornithonyssus (Macronyssidae), with inset
of chelicerae of Dermanyssus (Dermanyssidae); (C) hard tick, Amblyomma (Ixodidae). Original by Nathan D. Burkett-Cadena.
FIGURE 2.4 Legs of medically important insects. (A) Hind leg of blister
beetle, Epicauta (Meloidae); (B) hind leg of cat flea, Ctenocephalides felis
(Pulicidae); (C) foreleg of human body louse, Pediculus humanus (Ped-
iculidae); (D) hind leg of human pubic louse, Pthirus pubis (Pthiridae).
Original by Nathan D. Burkett-Cadena.
flight mechanism in flying insects, helps to store energy and
significantly increase the efficiency of the hind legs in the
jumping ability of fleas. Contraction of muscles in the hind
legs compresses pads of resilin located at the bases of the
hind coxae. A release mechanism causes rapid expansion of
the resilin pads, propelling the flea forward and upward
during the jump.
Lice are noted for their ability to cling tenaciously to
their hosts. The legs of sucking lice are particularly well
adapted for grasping host hair. The claws are formed by
modifications of the tibia and tarsus, called tibiotarsal claws
(Fig. 2.4C and D). The tarsus is generally reduced to form a
stout, curved, often sickle-shaped, movable element. This
element articulates with a sclerotized projection of the tibia
to form an effective grasping structure.
Among the Diptera, hippoboscids (e.g., keds) have legs
that are specialized for host attachment. Unlike other blood-
feeding dipterans, for which host contact is usually brief,
hippoboscids may spend their entire lives on their hosts.
The legs are therefore stout and usually spinose, with
enlarged tarsal claws. These features enable hippoboscids
to hold onto their host and to move about quickly and
efficiently amid host pelage.
In mites, each leg typically consists of seven segments
(Fig. 2.5A). The basal segment is the coxa, followed by the
trochanter, femur, genu, tibia, tarsus, and terminal apotele.
The apotele may bear claws, setae, and/or an empodium, a
padlike structure arising between the bases of the claws. In
parasitic mites, various modifications of the legs enable
 Morphological Adaptations of Parasitic Arthropods Chapter | 2
FIGURE 2.5 Legs of medically important mites. (A) Leg I of Pneu-
molaelaps (Laelapidae); (B) leg 1 of hard tick, Amblyomma (Ixodidae);
(C) leg 3 of fur mite Trichoecius (Myocoptidae); (D) leg I of scabies mite,
Sarcoptes (Sarcoptidae). Original by Nathan D. Burkett-Cadena.
these arthropods to locate and attach to their hosts. In ticks,
for example, the forelegs have enlarged claws (Fig. 2.5B)
that enable them to quickly grasp passing hosts and facil-
itate holding onto host skin or pelage during feeding and
mating. Certain feather mites (e.g., Analgidae) have sucker-
like empodia and large spurs on their legs for securing
themselves to their avian hosts. Many of the mites that
parasitize snakes (e.g., Ixodorhynchidae and Ophiony-
ssidae) also have sucker-like empodia, which facilitate
movement on their host and holding onto the smooth sur-
faces of the body scales. In many other parasitic mites, hind
legs, rather than forelegs, are modified for host attachment.
Mites in the family Mycoptidae, for example, have legs 3
and 4 enlarged with opposable digits for clasping the fur of
their mammalian hosts (Fig. 2.5C). In scabies mites (Sar-
coptidae), all four pairs of legs are reduced and have
21
elongate apoteles with terminal suckers (Fig. 2.5D). These
structures allow the mites to move about quickly on the
surface of the skin and to hold tightly to the epidermis.
SENSORY STRUCTURES
Sensory structures play an integral role in host location and
recognition and thus have become highly specialized in
many parasitic arthropods. Various sensory structures of
parasitic arthropods function to detect motion, vibrations,
temperature, moisture, carbon dioxide, and a plethora of
chemical substances produced by potential hosts. In com-
bination with each other, these environmental and host-
associated cues are often specific for a single host species
or group of closely related host animals.
In insects, the antennae and eyes are the primary sensory
organs. The antennae of blood-feeding insects, particularly
hematophagous dipterans, have receptors that detect chem-
icals emanating from the skin and present in the exhaled
breath of potential hosts. Host substances known to attract
mosquitoes include carbon dioxide, lactic acid, octenol, es-
trogen, fatty acids, and amino acids. In mosquitoes, sensory
receptors in the basal segment of the antenna are highly
developed to form the Johnston’s organ, which is special-
ized for detecting airborne vibrations. Host-seeking female
mosquitoes may cue in on vibrations produced by host
movements and even vocalizations by hosts such as birds
and frogs. Fleas exhibit an interesting modification of the
antennae, in which the antenna is short, flattened, and fits
into a protective groove on the side of the head (Fig. 2.2D).
This allows the antennae to be retracted so as not to become
damaged or impede movement as the flea maneuvers amid
host hair or feathers.
In fleas and lice, the eyes are generally reduced in size
(Fig. 2.2D and F) or may be altogether absent. In some
cases, such modifications of the eyes help to prevent
damage to the sense organs, while in other cases the
reduction of eyes reflects the relative unimportance of
vision in the life of the parasite. In many other insects, such
as mosquitoes (Fig. 2.2B), biting midges (Fig. 2.2C), and
horse flies, the eyes are greatly enlarged, reflecting the more
significant role that light perception and vision play in
locating, or orienting toward, potential host animals.
Sensory structures are often particularly numerous or
well developed on the mouthparts of parasitic arthropods.
In many solenophages, for example, these receptors are
concentrated near the tip of the proboscis or rostrum
(e.g., mosquitoes and bed bugs, respectively) and are used
to detect the precise location of capillaries beneath the
surface of the skin. In biting midges, sensory receptors for
detecting environmental and host cues are concentrated in a
specialized pit, in the form of a sensory organ on the
enlarged third segment of the maxillary palp (Fig. 2.2C).
22 Medical and Veterinary Entomology
In fleas, the dorsal portions of the terminal abdominal
segments are modified as a sensory organ, called the sen-
silium. The associated sensory structures are specialized for
detecting host-associated cues such as vibrations and tem-
perature gradients.
In mites, chemical and tactile cues are perceived by
sensory structures on the pedipalps, legs, and various other
parts of the body. Specialized sensory setae with an asso-
ciated socket-like base, called trichobothria, are common
in many groups of mites and other arachnids for detecting
airborne and substrate vibrations and other tactile cues. In
certain groups of mites, the first pair of legs may be un-
usually long and slender, with numerous receptors, serving
as a sensory organ, much like the antennae of insects. In
ticks, a complex sensory structure, called Haller’s organ,
is located on the dorsal aspect of the tarsus of the first pair
of legs and functions in detection of temperature, air
movements, host odors, and other host and environmental
cues.
In summary, a number of advantageous morphological
modifications are evident in ectoparasitic and blood-feeding
arthropods. Recurring adaptations include modifications of
the body shape, feeding apparatus, locomotory appendages,
and sensory structures. Each modification has allowed
parasitic arthropods to more efficiently exploit their verte-
brate hosts. A number of excellent works have been pro-
duced, such as those by B. Jobling (1987) and R. E.
Snodgrass, that examine in detail the morphological adap-
tations of parasitic arthropods. For further information on
general insect and arachnid morphology, see Snodgrass
(1993), Chapman (1998), and Krantz and Walter (2009).
REFERENCES AND FURTHER READING
Chapman, R. F. (1998). The insects: Structure and function (4th ed.).
Cambridge: Cambridge University Press.
Harbach, R. E., & Knight, K. L. (1980). Taxonomist’s glossary of mos-
quito anatomy. Biological Research Institute of America, Plexus
Publishing, Inc.
Hopkins, G. H., & Rothschild, M. (1953). An illustrated catalogue of the
Rothschild collection of fleas (Siphonaptera) in the British Museum
(Natural History) (Vol. I). London: Tungidae and Pulicidae, British
Museum (Natural History).
Jobling, B. (1987). Anatomical drawings of biting flies. London: British
Museum (Natural History).
Krantz, G. W., & Walter, D. E. (Eds.). (2009). A manual of acarology (3rd
ed.). Texas Tech University Press.
Labrzycka, A. (2006). A perfect clasp - adaptation of mites to parasitize
mammalian fur. Biological Letters, 43, 109e118.
Lozano-Fernandez, J., Carton, R., Tanner, A. R., Puttick, M. N.,
Blaxter, M., Vinther, J., et al. (2016). A molecular palaeobiological
exploration of arthropod terrestrialization. Philosophical Transactions
of the Royal Society B, 371(1699), 20150133.
Marshall, A. G. (1981). The ecology of ectoparasitic insects. London:
Academic Press.
Regier, J. C., Shultz, J. W., Zwick, A., Hussey, A., Ball, B., Wetzer, R.,
et al. (2010). Arthropod relationships revealed by phylogenomic
analysis of nuclear protein-coding sequences. Nature, 463,
1079e1083.
Sasaki, G., Ishiwata, K., Machida, R., Miyata, T., & Su, Z. H. (2013).
Molecular phylogenetic analyses support the monophyly of Hexapoda
and suggest the paraphyly of Entognatha. BMC Evolutionary Biology,
13, 236.
Snodgrass, R. E. (1943). The feeding apparatus of the biting and disease-
carrying flies: A wartime contribution to medical entomology.
Smithsonian Miscellaneous Collections, 104, 1e51.
Snodgrass, R. E. (1944). The feeding apparatus of the biting and sucking
insects affecting man and animals. Smithsonian Miscellaneous Col-
lections, 104, 1e113.
Snodgrass, R. E. (1946). The skeletal anatomy of fleas (Siphonaptera).
Smithsonian Miscellaneous Collections, 104, 1e110.
Snodgrass, R. E. (1993). Principles of insect morphology. Ithaca, NY:
Cornell University Press.
Stojanovich, C. J. (1945). The head and mouthparts of the sucking lice
(Insecta: Anoplura). Microentomology, 10, 1e46.