ENTOMOLOGY 322 LAB 2

Introduction to Arthropoda

The phylum Arthropoda (to which insects belong) is a huge and morphologically diverse group of over
1 million described species and almost certainly up to 20 to 50 million species in total. The arthropods arose
sometime in the early Cambrian period (>560 mya), during what is called the Cambrian explosion, when the
first multicellular life appeared in the fossil record.
The phylogenetic relationships among the arthropod classes has remained a subject of debate in sys-
tematics. While morphological studies have generally supported the cladogram in Fig. 2.1 (adapted from
Brusca & Brusca, 1990; Weygoldt, 1986 and Wheeler et al. 1993), in which Hexapoda is sister to Myriapoda,
many molecular studies (summarized in Fig. 2.2 a-f) have supported a sister group relationship between the
Hexapoda and the Crustacea. The most recent and definitive molecular study is that of Shultz & Regier (2000)
in which they analyzed two slowly-evolving nuclear genes (EF-1alpha and Pol II) based on parsimony and
maximum likelihood. Their results provide strong support for the hypothesis that Hexapoda are the sister group
to Crustacea (Fig. 2.3b) or arise from within the Crustacea (Fig. 2.3a). These results corroborate earlier
studies by the same authors (Regier & Shultz 1997, 1998). These new molecular studies are extremely exciting
because they suggest that hexapods are terrestrial crabs! Keep in mind, however, that there remains much
controversy about the true affinities of the hexapods.

Hexapoda
(insects and relatives)

Figure 2.1
 (a)
(d)

(b)
(e)

(c)
(f)

Figure 2.2 (Note: "Myriopods" should be "Myriapods.")

Figure 2.3 Phylogeny of the Arthropoda based on combined
EF-1alpha and PolII for 17 taxa of arthropods (from Shultz &
Regier 2000). 2.3a shows the parsimony analysis of the
protein sequences. 2.3b shows the maximum likelihood
analysis of nucleotides. Note that the Hexapoda arise from
within the Crustacea or are sister to the Crustacea. There is no
evidence of Hexapoda + Myriapoda (=Atelocerata). Schultz &
Regier term the Hexapoda + Crustacea the "Pancrustacea."
 Figure 2.4 Cephalic appendages of Peripatus (Brusca & Brusca, Figure 2.5 Transverse section of Peripatopsis
1990) (Manton, 1977)

1.
Examine the demonstration of an Onychophora. Onychophora are unusual organisms found in
the tropics that share features with both the Annelids examined in lab 1 and the arthropods. Because of
this, the Onychophora has been extremely difficult to place phylogenetically. They share several derived
features (synapomorphies) with arthropods, such as (a.) growth by ecdysis and (b.) a greatly reduced
coelom, while also possessing primitive, annelid-like features (symplesiomorphies), such as (c.) paired
nephridia in each segment of the body, (d.) weak sclerotization, and (e.) unjointed appendages. Some
authors go so far as to place the Onychophora well inside the Arthropoda as the sister group to the
Myriapods+Insects. This seems an unlikely hypothesis based on the mix of characteristics mentioned above
(for example, paired nephridia would have had to arise independently in Annelids and Onychophora).
Use the accompanying diagrams to locate the antennae, the jaws, and the oral papillae or slime
papillae. The oral papillae are used to project a sticky, slime-like material for capturing their arthropod
prey. Examine the body appendages or
lobopods. Are the anterior lobopods similar to
the posterior ones?
The cuticle of Onychophora is covered
in tiny tubercles. The microsculpture of these
is useful in species-level identification. Do you
see any sclerites? How might this be related to
their being limited to tropical regions?

2.
Next, we will observe a member of the
likely sister group to the Arthropoda:
Tardigrada. Tardigrades also share a number
of derived traits with the Arthropoda not Figure 2.6 Styraconyx quivitoq, ventral and dorsal views
possessed by the Annelida (hence- their (Kristensen & Higgins, 1984)
 Figure 2.7 Internal morphology of Tardigrada Figure 2.8
(Cuenot, 1949)

placement in the phylogeny closer to Arthropoda than Annelida). These traits include (a.) the evolution
of discrete, segmental, striated muscle bands attached to apodemes, (b.) shift from lobopodal locomotion
to true leg-gait movement, (c.) loss of serially arranged nephridia (remember, these were present in
Onychorphora), (d.) elongate, cuticular setae, and (e.) segmental plates over much of the body which
may be homologous to arthropod sclerites.
Observe the demonstration of live Tardigrades. Observe their locomotion -- can you tell where
the leg muscles originate and insert? How do they control the movements of their legs? Would you
describe their mode of locomotion as arthropod-like or annelid-like? Can you tell how they feed? Do
you detect any internal segmentation of the body (such as intersegmental septae, serially arranged
metanephridia, or serially arranged ganglia? Tardigrades lack respiratory and circulatory systems -- can
you propose an explanation for this?

3.
Next, we shall examine a representative arthropod, the centipede Scolopendra viridis (a member
of the Class Myriapoda, the presumed sister group to insects). Our purpose is to contrast the basic body plan
of an arthropod with that of an annelid.

The phylum Arthropoda is united by a number of unique and unreversed characteristics

(synapomorphies):

a. the presence of paired compound eyes and one to several median simple eyes.
b. rigid, articulated exoskeleton composed of cuticle, which is hardened through calcification (mineral
deposition) or sclerotization (protein cross-linking). Exoskeleton typically consists of dorsal
tergites, lateral pleurites and ventral sternites although fusion to various degrees is common.
c. external and internal body segmentation, with strong tendency toward regional specialization and
tagmosis.
d. articulated, jointed appendages located on each body segment; appendages with intrinsic, striated
musculature typically with antagonistic (paired flexor and extensor) muscles.
e. coelom reduced to portions of reproductive and excretory system -- main body cavity is an open
hemocoel.
f. appendages positioned ventrally.
g. loss of all motile cilia and flagella except in sperm.
 Obtain a preserved centipede and place it in 70% ethanol in a dissecting pan. The specimen
may be handled dry when examined under the microscope, but do not allow it to dry out. Note that
like annelids, onychophorans and tardigrades, the centipede is obviously segmented when viewed
externally. However, unlike these groups, the body is divided into two discrete regions (tagmata), the
head and the trunk.
Tagmosis (character c, above) is an important feature which distinguishes arthropods from
non-arthropod relatives. Tagmosis refers to regional specialization of body segments. Neighboring
segments become closely associated and partially fused to form a discreet group of segments, a tagma
(plural - tagmata). Each tagma differs in overall appearance and function from other tagmata. In
Chelicerata and Pycnogonida the body is divided into two tagmata: cephalothorax and abdomen. In
insects and crustaceans the body is divided into three tagmata: head, thorax and abdomen. [If available,
briefly examine a scorpion or a horseshoe crab]

Figure 2.9 Diagramatic representation of arthropod tagmosis (Snodgrass, 1935)

 Figure 2.11 Maxilliped (dorsal) showing poison gland
outlet

Figure 2.10 Lithobius forficatus (Rilling, 1968)

The trunk segments alternate between longer and shorter segments. Each segment of the trunk
bears a pair of legs, serially homologous with each other (character d, above). How do the legs of a
centipede (or any other arthropod) differ from the legs of an Onychophoran in terms of musculature?
On close examination, the legs are not all identical. The posterior legs are enlarged and are used as
defensive pinchers when attacked from the rear.
The anterior pair of trunk appendages are thick, heavily sclerotized maxillipeds (Fig. 2.10, Mxp) that
underlie the head and appear to be part of it when viewed ventrally. The maxillipeds are in fact not part of the
head, but are homologous to the first pair of thoracic legs in insects, the prothoracic legs. The maxillipeds
contain poison glands, end in sharp points, and are used in prey capture and in defense. Centipedes can inflict
a painful bite!
The head externally is a sclerotized capsule in which intersegmental grooves are absent. The presence
of multiple segments can be detected externally by the appendages, which are serially homologous with the
trunk legs, although of quite different form. A pair of long sensory antennae project anteriorly. Concealed by
the maxillipeds are three pairs of feeding appendages (Fig. 2.10), the mandibles, first maxillae (Mx1) and
second maxillae (Mx2). These have been exposed in a demonstration. Note also a cluster of four simple eyes
(character a, above) just posterior to each antenna.
Examine a long trunk segment near the caudal end that has a spiracle. Note that the integument
is composed of hard sclerites (character b, above) and softer, more flexible membranes and that a sclerite
need not be melanized (darkly pigmented) to be hard. Probe with forceps or pin-point to note the
difference between sclerite and membrane. Each segment has a dorsal sclerite, the tergum, and a ventral
sclerite, the sternum. Laterally the pleuron is largely membranous except for where the leg articulates
with the body. Observe the pleural region and identify the subcoxal sclerites (Fig. 2.12E, scx). How many are
there and what are their positions relative to the leg (dorsal, ventral, anterior, posterior?). The spiracle, the
Figure 2.12 Otocrytops sexspinosa (Snodgrass, 1952)
opening to the tracheal system, is in the pleural membrane above the leg (Fig 2.13). Does every segment have
a spiracle?
Gently move the leg and note how it articulates with the body (Fig 2.14). The coxa articulates with the
overlapping sternum. The weakly sclerotized area in front of the coxa is the procoxa (=subcoxa). The
procoxa is more heavily sclerotized in the more posterior segments. It is not a separate leg segment (=podite)
and probably represents sclerotization of the pleuron [Note: the procoxa is probably homologous to the insect
pleural sclerite or pleuron]. The trochanter is a narrow podite that articulates with the coxa. The more distal
podites are the prefemur (=2nd trochanter), femur, tibia, 2-segmented tarsus, and small pretarsus, represented
only by the terminal claw. (Note that the tarsus is one-segmented on the accompanying Fig 2.12 of Otocrytops.)
Cut open one leg and identify the internal striated muscles that allow each leg segment to move independently
(character d, above).

Figure 2.13 Lithobius. Limb morphology (Manton,

1965)

Figure 2.14 Lithobius sp. lateral aspect showing spiracle

4.
Now lets turn to insects and examine the insect exoskeleton as a whole, in order to identify
general “design features” of this skeleton. Obtain a pickled specimen of the lubber grasshopper, Romalea
micropteryx (rinse it in water to wash off the preservative). Under the microscope examine the surface
of the body and note that some tagmata are more heavily armored than others. (You may need to poke
the exoskeleton with a pin to convince yourself of this.) Which tagma is most heavily armored (sclerotized)
and why; which is the least heavily armored?
Now focus your attention on the abdomen. Note that, as in the centipede, individual sclerites
form the top and bottom of each segment, and that between these sclerites are flexible regions composed
of arthrodial membrane. What developmental process causes the distinction between membrane and
sclerite in arthropods? Move the abdomen from side to side and note how each segment moves with
respect to its neighboring segments. The abdomen of insects retains several primitive arthropod features,
such as homonomy (similarity among each segment with little differentiation in either form or function;
as in the centipede), but also shows several derived features as well, such as the lack of jointed appendages
on each segment.
Now focus on the thorax. In general, how does it compare to the abdomen? Is it more or less
heavily sclerotized? Where are the arthrodial membranes located? Are individual segments of the thorax
flexible or tightly fused together? How does the insect abdomen compare to a group of three segments
in the centipede trunk?
Note that in lateral view the side of the thorax is marked by distinct lines. Some of these lines
(called sutures) represent the intersegmental boundaries. Others (called sulci; sulcus=singular) represent
internal ridges (called phragmata; phragma=singular). Phragmata are structural elements of the insect
exoskeleton which perform two important functions: (1) they are a lightweight means of providing
strength to the exoskeleton by bracing its inner walls, and (2) they provide sites for muscle attachment.
We will be examining sulci and their associated phragmata in great detail in future labs; but for now,
observe the cleared thorax of Romalea and note the complexity of the internal skeletal anatomy. Insect
skeletons are not simply pill boxes enclosing the insect in a hard case -- they are complex structures with
intricate connections to the internal muscles and tendons!
Now briefly examine the head of your grasshopper. Note that it is also heavily sclerotized, like
the thorax. The lines on the head capsule correspond to a mixture of true sutures, sulci and a third type
of line, an ecdysial cleavage line. These are typically fainter than sulci and correspond to the lines along
which the exoskeleton splits during molting. One ecdysial cleavage line in Romalea runs down the
midline of the head capsule in dorsal view.

Figure 2.15 (Daly et al. 1987)

As vertebrate animals we have a tendency to view internal skeletons as the best type of skeleton
around. Internal skeletons do in fact have some beneficial attributes, such as allowing gradual growth
throughout life. However, from a purely architectural perspective, arthropod exoskeletons have many
advantageous features. First, they are made of chitin, a long-chain polysaccharide, embedded within a
matrix of proteins. This combination of chitin and protein (cuticle) is extremely strong for its weight (in
fact stonger than bone) and is flexible, unlike bone, so it can withstand bending forces with less damage.
Secondly, the arrangement of the skeleton outside of the muscle is a more efficient way to allocate
skeletal material. By concentrating skeletal material toward the outside of an animal, the skeleton has
greater resistance to bending forces. To illustrate this point, think about the relative ease in bending a
hollow tube vs. a rod of equal mass. While external skeletons may place some upper limits on arthropod
body size, for small animals, they are ideal.