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species of Trilobites belonging to 39 genera ranging from Lower
Cambrian to Carboniferous.
A “metastoma” or lower lip plate (Fig. 142, Ep) is found just
behind the hypostome in Triarthrus, but has not been noticed in any
other genus. Between the hypostome and the metastoma lies the
mouth.
The segments of the thorax are free, and their number varies
from two in Agnostus (Fig. 146) to twenty-six in Harpes (Fig. 150,
A). In the Trilobites confined to the Cambrian period the number
(except in the Agnostidae) is usually larger than in the genera found
in the Ordovician and later periods. Owing to the free thoracic
segments many Trilobites were able to curl up somewhat after the
manner of a Wood-louse (Figs. 137, D, 138). The axial part of each
thoracic segment is more or less considerably arched. Usually it
consists of three parts: (i.) the largest part (Fig. 137, C, a), called the
ring, is band-like in form, and is always visible whether the Trilobite
is extended or coiled up; (ii.) in front of the ring is a depressed,
groove-like part (Fig. 137, C, b) separating it from (iii.) the articular
portion (c) which is convex in front and extends beneath the ring of
the preceding segment; this part is only visible when the Trilobite is
coiled up or when the segments are separated. In some few genera
the axial part consists of a simple arched band without either a
groove or a specially modified articular portion. The pleurae (Fig.
137, A, l, C, d-f) are fixed firmly to the axis, and have the form of
narrow bands with the ends rounded, obtuse, pointed, or spinose. In
a few cases the pleurae have a plain surface; but usually they possess
either a ridge or a groove (Fig. 137, C, g); the former is generally
parallel to the margins of the pleura, the latter is generally oblique,
being inclined backwards from the axis. Sometimes in front of the
ridge there is a small groove. On the ventral surface each pleura
shows, at its outer extremity, a reflexed margin or doublure. At some
distance from the axis the pleurae are bent downwards and
backwards. The point where this bend occurs is called the “fulcrum”
(e); it divides the pleura into an internal and an external part: the
internal part (d-e) is flat or slightly convex, and just touches the front
and back margins of the adjacent pleurae; the external part (e-f) may
be (i.) narrower than the internal part, so that it is separated from
the previous and succeeding pleurae; such occurs principally in
pleurae with ridges, as in Cheirurus and Bronteus; or (ii.) it may be
in the form of a long cylindrical process, as in many species of
Acidaspis; or (iii.) the external part may be of the same width, either
throughout or in part, as the internal part, and may overlap the next
pleura behind; this type is found principally in pleurae with a groove
such as in Phacops, Calymene, Sao, Asaphus, Ellipsocephalus.
In some Trilobites there is beyond the fulcrum a smooth, flat,
triangular part at the front margin of the pleura; this part is known
as the “facet,” and forms a surface articulating with the preceding
segment which overlaps it.
In the remarkable form Deiphon (Fig. 151, E) the pleurae are
separate throughout their entire length.
In some Trilobites broad and narrow forms of the same species
occur—the difference being seen especially in the axis. The former
are regarded as females, the latter as males.[190]
The segments of the abdomen or pygidium (Fig. 137, A, 3) are
similar to those of the thorax, except that they are fused together. In
a few forms, such as Illaenus (Fig. 150, F) and Bumastus, the fusion
is so complete that no trace of segmentation can be seen on the
dorsal surface. Usually, however, the segments are easily
distinguishable; the number seen on the axis is commonly greater
than on the lateral parts of the pygidium; this difference is
particularly well shown in Encrinurus. In Trilobites which have
grooved pleurae the conspicuous grooves seen on the lateral parts of
the pygidium are the grooves of the pleurae, the sutures between the
pleurae being less distinct. The shape of the pygidium may be
semicircular, a segment of a circle, trapezoidal, triangular, semi-
parabolic, etc.; its size varies considerably; in the Cambrian forms it
is usually small, but in the Trilobites of later periods it becomes
relatively larger. The number of segments in the pygidium varies
from two to twenty-eight. The axis of the pygidium tapers more
rapidly than that of the thorax; sometimes it reaches quite to the
posterior end of the body, but is commonly shorter than the
pygidium; in Bronteus it is extremely short, and the grooves on the
lateral parts of the pygidium radiate from it in a fan-like manner.
Occasionally, as in Bumastus, the axis cannot be distinguished from
the lateral parts. In a few early Trilobites (Olenellus, Holmia, Fig.
148, Paradoxides, Fig. 147) the lateral parts of the pygidium are very
small. In some genera, such as Asaphus, the marginal part of the
pygidium forms a flattened or concave border. The margin may be
entire or produced into spines, and sometimes (Fig. 151, C) a caudal
spine comes off from the end of the axis. On the ventral surface of
the pygidium there is a marginal rim similar to the doublure of the
cephalic shield. The anus is on the ventral surface of the last segment
of the pygidium.
Although Trilobites are often found in abundance and in an
excellent state of preservation, it is only in very rare cases that
anything is seen of the ventral surface except the hypostome and the
reflexed borders of the cephalic shield, of the thoracic segments, and
of the pygidium. The usual absence of appendages is probably due
to their tenuity. Billings, in 1870, first obtained clear evidence of the
presence of pairs of appendages, in Asaphus platycephalus. Soon
afterwards Walcott[191] showed their existence in American
specimens of Asaphus megistos, Calymene senaria, and Cheirurus
pleurexacanthus. In the two latter species the appendages were
found by cutting sections of curled-up specimens obtained from the
Trenton Limestone; 2200 examples were sliced, of which 270
showed evidence of the existence of appendages. They were seen to
be present on the head, thorax, and pygidium; a ventral uncalcified
cuticle with transverse arches was also found. By means of sections
of curled-up specimens it was difficult to determine satisfactorily the
form and position of the appendages. Subsequently extended
specimens of Triarthrus (Fig. 142) and Trinucleus, showing the
ventral surface and appendages clearly, were discovered in the Utica
Slate (Ordovician) near Rome, New York. A full account of the
appendages in those specimens has been given by Beecher.[192]
In Triarthrus each segment, except the anal, bears a pair of
appendages, all of which, except the first, are biramous. There are
five pairs of cephalic appendages; the first pair are attached at each
side of the hypostome, and have the structure of antennae, each
consisting of a single flagellum formed of short conical joints. The
other cephalic appendages increase in size successively. At present
the second and third pairs are not satisfactorily known, but appear to
have been similar to the fourth and fifth pairs. The second pair is
attached at the level of the posterior end of the hypostome. The
fourth and fifth pairs have large, triangular coxopodites which served
as gnathobases, their inner edges
being denticulate; the
endopodites consist of stout
joints; the exopodites are slender,
and bear setae which are often
arranged in a fan-like manner.
The first pair of appendages
appear to be antennules, whilst
the second pair probably
represent the antennae, the third
pair the mandibles, and the
fourth and fifth pairs the maxillae
of other Crustacea. The
appendages of the thorax and
pygidium do not differ essentially
from the two posterior cephalic
appendages. Those on the
anterior part of the thorax are the
longest; the others gradually
decrease in size in passing
posteriorly. Each thoracic leg
(Fig. 142, B) consists of a short
coxopodite with an inward
cylindrical prolongation forming
a gnathobase which is best
developed on the anterior legs;
the endopodite and exopodite are
long and nearly equal; the former
consists of six joints tapering
gradually to the end; the latter, of Fig. 142.—Triarthrus becki, Green, ×
a long proximal joint with a 2½. Utica Slate (Ordovician), near
denticulate edge and a distal part Rome, New York. A, Ventral surface
of ten or more joints, and it bears with appendages; Ep, metastome; Hy,
hypostome. B, second thoracic
a row of setae along the whole of appendage; en, endopodite; ex,
the posterior edge. exopodite, × 12. (After Beecher.)
The anterior appendages of the
pygidium differ but little from the
posterior thoracic legs; but the phyllopodous character, which
appears in the latter, becomes more distinct in the appendages of the
pygidium, especially those near its posterior end, and is due to the
broad, flat, laminar joints of the endopodite.
The more striking features of the appendages of Triarthrus are the
small amount of differentiation which they show in different parts of
the body, especially the want of specialisation in the cephalic region;
the distinctly biramous character of all except the first pair; and the
presence of one pair of functional antennae only, and the occurrence
of thoracic gnathobases.
In Trinucleus the appendages are not so well known, but they are
considerably shorter than in Triarthrus.
In the Palaeozoic rocks of Bohemia, where Trilobites are very
perfectly preserved, Barrande[193] discovered the larval forms of
several species, and in some cases was able to trace out the
development very completely, but in others the earliest stages were
not found. In the strata in which Trilobites occur Barrande found
minute spheroidal bodies, usually of a black colour, and only a little
smaller than the youngest larval stages; those bodies are probably
the eggs of Trilobites. Since the publication of Barrande’s work the
development of some species found in North America has been
studied by Ford, Matthew, Walcott, and Beecher. But even now the
development is known in only a very small proportion of the total
number of genera of Trilobites. The early larval form (Fig. 143, A) is
similar in general character in the various species in which it has
been found. It is circular or ovoid in outline, with a length of from
0·4 to 1 mm., and consists of a large cephalic and a small pygidial
portion; the axis is distinct and usually shows more or less clear
indications of five cephalic segments; the eyes, when present, are
found at or near the front margin, and the free cheeks, if visible at all
on the dorsal surface, are narrow. For this early larval form Beecher
has proposed the name “protaspis”; he regards it as the
representative of the Nauplius of other Crustacea, but that view is
not accepted by Professor J. S. Kingsley.[194]
The general changes which occur in the course of development are:
modifications in the shape and relative size of the glabella, and of the
number and depth of the glabella-furrows; the growth of the free
cheeks and the consequent inward movement of the facial sutures
and eyes; the introduction of and gradual increase in number of the
thoracic segments, and the relative decrease in size of the head.
Sao hirsuta is a species found
in the Cambrian, the
development of which was fully
described by Barrande. Its
earliest protaspis stage (Fig. 143,
A) is circular in outline; the
glabella expands in front and
reaches the anterior margin; the
pygidial region is not distinctly
separated from the cephalic
Fig. 143.—Development of Sao hirsuta,
Barr. Cambrian. A, Protaspis; B-F,
region; segmentation is indicated
later stages; G, adult. The small in the former, and the neck-ring
outlines below each figure show the is present in the latter; the eye-
actual size of each specimen. (After line is seen on each side of the
Barrande.) glabella near the anterior margin.
In a later stage (Fig. 143, C) the
segmentation of the glabella becomes more distinct, indicating the
existence of five cephalic segments, and the facial suture appears
near the margin limiting a very narrow free cheek. Subsequently
(Fig. 143, D-F) the thoracic segments develop, and increase in
number until the adult stage (G) is reached; also the eyes appear at
the margin of the cephalic shield, and gradually move inwards, and
the glabella becomes narrower and rounded in front, and ceases to
reach the anterior margin. In this species the eye-line is present in
the adult.
In the protaspis of Triarthrus (Fig. 144), found in the Ordovician,
the glabella does not reach the front margin nor expand in front as it
does in Sao; an eye-line is present, but disappears before the adult
stage is reached.
 Fig. 144.—Triarthrus becki, Green.
Ordovician. A, B, Two successive stages
of the protaspis, × 45. (After Beecher.)

Fig. 145.—Larval stages of Trilobites. A-D, Dalmanites socialis,

Barr. Ordovician, Bohemia. The small figures below show the
natural size of each specimen. (After Barrande.) E, Mesonacis
asaphoides, Emmons, × 10. Lower Cambrian, North America.
(After Walcott.) F, Acidaspis tuberculata, Conrad, × 20. Lower
Helderberg Group (Lower Devonian or Upper Silurian), Albany
County. (After Beecher.)
 Dalmanites (Fig. 151, C) is a more advanced type than Sao and
Triarthrus, and is found in later deposits. In the earliest stage (Fig.
145, A) the head and pygidium are quite distinct, and there is no eye-
line present at this or any stage in development, but large ovoid eyes
are found on the front margin, and have their long axes placed
transversely to the axis of the body; the glabella is strongly
segmented and is rounded in front. In later stages (C, D) the
pygidium increases in size relatively, and the thoracic segments are
successively introduced; the facial sutures and free cheeks appear on
the dorsal surface, and as the free cheeks grow the eyes move
inwards and backwards, and gradually swing round until their long
axes become parallel with the axis of the body.
The larval form of Acidaspis (Fig. 145, F) is of interest since even
in the earliest stage it shows the spiny character which forms such a
striking feature of the adult (Fig. 151, F).
Before the discovery of the ventral surface of Trilobites it was
thought by some zoologists that their affinities were with the
Xiphosura rather than with the Crustacea. But the presence of
antennae, and of five pairs of cephalic appendages; the biramous
thoracic and pygidial appendages, the hypostome, and the character
of the larval form, as well as the absence of a genital operculum,
separate the Trilobites from the Xiphosura and connect them with
the Crustacea.
The position of the Trilobites in the Crustacea is, however, difficult
to determine. Already in the Cambrian period, at least five main
groups of the Crustacea were clearly differentiated, namely, the
Phyllopoda, Ostracoda, Cirripedia, Trilobita, and Leptostraca
(Phyllocarida), and probably also the Copepoda, but of the last no
remains have been preserved as fossils. Palaeontology, therefore,
furnishes no connecting links between any two of these orders.
The Crustacea to which the Trilobites show some resemblance are
the families Apodidae and Branchipodidae of the Order Phyllopoda
(see pp. 19–36). The Trilobita agree with those families in having a
large but variable number of trunk-segments, in the possession of a
large labrum (hypostome), and in the occurrence of gnathobases on
the thoracic appendages; also the foliation of some of the trunk-
appendages is somewhat similar. The points of difference, however,
are considerable; thus the cephalic appendages are much more
specialised in the Apodidae and Branchipodidae than in the
Trilobita; in the latter all, with the exception of the antennae, are
distinctly biramous, and whilst the basal joints were masticatory the
distal parts appear to have been locomotor organs. The appendages
of the trunk also differ considerably; in the Trilobita all are clearly
biramous, those of the thorax having a schizopodal form. In the
possession of a single pair of antennae the Trilobita differ from other
Crustacea; but in some forms of Apus the second pair of antennae
may be rudimentary or even absent.
There are still other features which characterise the Trilobita: thus
the eyes are borne on free cheeks, and differ in structure from those
of Phyllopods. The broad pygidium formed of fused segments and
without terminal fulcra is quite unlike the slender-jointed abdomen
of Apus and Branchipus; and whilst in the Trilobites all the segments
bear appendages, in the Phyllopods some, at any rate, of the
posterior segments are devoid of appendages. The distinct division of
the body into an axial and pleural region is not seen in Phyllopods,
and is probably a character of some importance, since it occurs in the
great majority of Trilobites, including all the early forms.
The existence of some relationship between the Trilobita and the
Leptostraca (Phyllocarida) has been maintained by Professor G. H.
Carpenter.[195] He points out that some of the earliest Trilobites, such
as Holmia kjerulfi (Fig. 148), possess nearly the same number of
segments as Nebalia (Fig. 76, p. 111), and that in the latter genus the
cephalic appendages, especially the mandibles and maxillae, are less
specialised than in Apus, and consequently differ less from those of
Trilobites than do the appendages of the Apodidae. Further, in
another genus of the Leptostraca, Paranebalia, the biramous
thoracic legs, in which both endopodite and exopodite are elongate,
approach those of Trilobites more nearly than do the thoracic legs of
Apus.
The view[196] that some connexion may exist between the Isopoda
and the Trilobita seems to have been based on the similar dorso-
ventral flattening of the body, its division into three regions—head,
thorax, and abdomen—and the presence of sessile eyes. Beyond this
it is difficult to find any resemblance; whilst the differences, such as
the variable number of thoracic segments and their biramous
appendages in Trilobites, are important.
 At present, then, we can only conclude that the Trilobita are more
primitive than any other Crustacea, and that their resemblance to
some of the Phyllopoda is sufficient to make it probable that they had
some ancestral connexion;[197] the possibility of such a relationship
receives some support from the presence in the Lower Cambrian
rocks of Protocaris, a genus of the Phyllopoda which resembles
Apus.[198] The primitive characters of Trilobites are the variable and
often large number of segments in the thorax and pygidium; the
presence of a pair of appendages on every segment except the anal;
the biramous form of all except the first pair of appendages; and the
lack of specialisation shown by the appendages, especially those of
the head.
The classification of Trilobites is due largely to the work of
Barrande and Salter, and the families defined by those authors have
been, in the main, generally adopted. But the phylogenetic
relationship of the families has still, to a large extent, to be
established. Salter[199] arranged the families in four groups, but did
not claim that that classification was entirely natural. His groups
with the families included in each are:—
1. Agnostini. Without eyes or facial suture. Agnostidae.
2. Ampycini. Facial sutures obscure, or submarginal, or absent.
Eyes often absent. Trinucleidae.
3. Asaphini. Facial sutures ending on the posterior margin.
Acidaspidae, Lichadidae, Harpedidae, Calymenidae, Paradoxidae,
Conocephalidae, Olenidae, Asaphidae, Bronteidae, and Proëtidae.
4. Phacopini. Facial sutures ending on the lateral margins. Eyes
well developed. Phacopidae, Cheiruridae, and Encrinuridae.
A modification of Salter’s classification has been brought forward
by Beecher[200] who divides the Trilobita into three main groups:—
1. Hypoparia. Facial sutures at or near the margin, or ventral.
Compound eyes absent. This is equivalent to Salter’s Agnostini and
Ampycini with the addition of the Harpedidae.
2. Opisthoparia. Facial sutures extending from the posterior
margin to the front margin, but occasionally uniting in front of the
glabella. Eyes holochroal or prismatic, but sometimes absent. This
comprises the same families as Salter’s Asaphini with the exclusion
of the Harpedidae and Calymenidae.
 3. Proparia. Facial sutures extending from the lateral margins,
and either cutting the anterior margin or uniting in front of the
glabella. Eyes holochroal or schizochroal; occasionally absent. This is
equivalent to Salter’s Phacopini with the addition of the
Calymenidae.
In each of the groups proposed Beecher regards as the more
primitive forms those which possess characters similar to those of
the early larval stages, such as narrow free cheeks, the absence of
compound eyes, and a glabella which is broad in front and reaches
the anterior margin of the head.
The modifications introduced by Beecher can scarcely be regarded
as making Salter’s classification more natural. For instance, the
Agnostidae differ so much from all other families that, at present,
there is no evidence to show that they have any close phylogenetic
relationship with the Trinucleidae and Harpedidae. Further, the
Calymenidae, which Salter recognised as related to the Olenidae,
have been shown by the careful work of Professor Pompeckj[201] to
have descended from the latter family, and to have no genetic
connexion with the Phacopidae with which they are grouped by
Beecher. Then in the Trinucleidae the earliest genus, Orometopus[202]
(Fig. 140, A), possesses compound eyes and facial sutures which
begin at the posterior margin and unite in front of the glabella; so
that, according to Beecher’s classification, that genus would belong
to the Opisthoparia, whereas the later genera (Trinucleus, etc.) of the
same family would be placed in the Hypoparia. At present, therefore,
the only classification of Trilobites which can be adopted is a division
into families, of which a short account is given below.
Fam. 1. Agnostidae (Fig. 146).—Small Trilobites, in which the
head and pygidium are of nearly the same size and shape. The thorax
is shorter than the head or pygidium, and consists of from two to
four segments with grooved pleurae. The length and width of the
head are commonly nearly equal, but sometimes the length is
greater. Eyes are absent. Facial sutures appear to be absent, but are
stated by Beecher to be at the margin of the cephalic shield. From the
absence of eyes, the probable absence of facial sutures, the few or
indistinct furrows on the glabella, and the smaller number of
thoracic segments, the Agnostidae appear to be degenerate forms.
Microdiscus is apparently less modified than Agnostus, on account
of the larger number of thoracic
segments, the more distinct
segmentation of the pygidium,
and, in some species, the larger
number of furrows on the
glabella. Cambrian and
Ordovician. Genera: Agnostus,
Microdiscus.
Fam. 2. Shumardiidae.—
The body is very small and oval.
The cephalic shield is nearly
semicircular and very convex,
with a broad glabella which
expands in front, and in which
the furrows, except the neck-
furrow, are indistinct. The facial
suture is marginal and eyes are
absent. There are six thoracic
segments with ridged pleurae; the
axis is broader than the pleurae.
The pygidium is large, and is
formed of about four segments
similar to those of the thorax.
Upper Cambrian and Ordovician.
Genus: Shumardia.
Fam. 3. Trinucleidae (Fig.
140).—The head is large and has a
flat border (except in Ampyx),
and long genal spines. In the
earliest genus (Orometopus) the Fig. 146.—Agnostus integer, Beyr., × 8.
facial sutures start from the Cambrian. (After Barrande.)
posterior margin (near the genal
angle) and pass obliquely inwards
to the compound eye, from whence they continue forward and unite
in front of the glabella. In Ampyx the suture starts from just within
the genal angle and passes to the front border, cutting off a narrow
free cheek; eyes are absent. In most specimens of Trinucleus no
sutures are seen, but some examples show indications of what may
be a facial suture (see p. 226), and a suture is sometimes found at the
margin of the cephalic border; eyes may occur (see p. 230). The
thorax consists of from five to eight segments, with grooved pleurae.
The pygidium is triangular. Principally Ordovician. Genera:
Orometopus (Upper Cambrian), Ampyx, Trinucleus, Dionide.
Fam. 4. Harpedidae (Figs. 139, G, H; 150, A).—The head is
large and has a broad, flat border which is finely punctate, and
extends backwards on each side in the form of a horn-like projection
nearly as far as the posterior end of the thorax. The glabella is convex
and does not reach the front margin. The cheeks are less convex than
the glabella, and bear eyes which usually consist of two or three
lenses. An eye-line connects the eye with the anterior part of the
glabella. A suture is stated to occur at the external margin of the flat
border. The thorax consists of from twenty-five to twenty-nine
segments; its axis is narrow, and the pleurae are long and grooved.
The pygidium is very small, and consists of three or four segments.
Ordovician to Devonian. Genus: Harpes.
Fam. 5. Paradoxidae (Figs. 147, 148, 149).—The cephalic shield
is large, and bears long genal spines. The glabella is more or less
swollen in front. The facial sutures appear to be absent in some
genera, and when present extend from the posterior to the anterior
margin. The palpebral lobes are long, and often more or less
semicircular or kidney-shaped. The thorax is long, and consists of
from sixteen to twenty segments with their pleurae produced into
spines. The pygidium is very small, and plate-like, or sometimes in
the form of a long spine. Cambrian. Genera: Olenellus, Holmia,
Mesonacis, Olenelloides, Paradoxides, Zacanthoides, Centropleura
(Anopolenus). Remopleurides (Fig. 150, D) from the Ordovician is
usually included in the Paradoxidae, but probably belongs to a
separate family.
Fam. 6. Conocephalidae (Conocoryphidae) (Fig. 150, E).—
The cephalic shield is semicircular, and larger than the pygidium.
The glabella narrows in front. The facial suture passes from near the
genal angle on the posterior border to the antero-lateral margin, and
limits a large fixed cheek and a narrow free cheek. Eyes are absent or
rudimentary, but an eye-line is usually present. The thorax consists
of from fourteen to seventeen segments with grooved pleurae, which
may be pointed, but are not usually produced into spines. The
 pygidium is small, and formed of
few segments. Cambrian. Genera:
Conocoryphe, Atops,
Ctenocephalus, Bathynotus.
Fam. 7. Olenidae (Figs. 142,
143; 150, B, C).—The cephalic
shield is larger than the
pygidium. The glabella is either
rectangular or parabolic. The
facial suture passes from the
posterior to the anterior margin.
The palpebral lobes are of
moderate or rather large size, and
are connected by an eye-line with
the front part of the glabella. The
thorax includes from eleven
(occasionally fewer) to eighteen
segments with grooved pleurae.
The pygidium is usually small,
with from two to eight segments.
Principally Cambrian. Genera:
Ptychoparia, Angelina,
Solenopleura, Sao, Agraulos
(Arionellus), Ellipsocephalus,
Protolenus, Olenus, Peltura,
Acerocare, Eurycare, Ctenopyge,
Fig. 147.—Paradoxides bohemicus, Leptoplastus, Triarthrus,
Barr. × ½. Middle Cambrian. (After
Zittel.) Parabolina, Sphaerophthalmus,
Parabolinella, Ceratopyge
(position doubtful).
Dikelocephalus is usually placed in the Olenidae, but perhaps
belongs to a distinct family.
Fam. 8. Calymenidae (Figs. 136, 137).—The glabella is broadest
behind. The facial suture starts at or near the genal angle—
sometimes on the posterior border just inside the angle, sometimes
on the lateral border just in front of the angle; the suture may be
continuous with the other suture in front of the glabella, or may cut
the anterior margin, beneath which it is connected with the other
suture by means of a transverse
suture (Fig. 137, B, D). The eyes
are rather small. The thorax
consists of thirteen segments with
grooved pleurae; the pygidium of
from six to fourteen segments.
Ordovician to Devonian. Genera:
Calymene, Synhomalonotus,
Homalonotus.

Fig. 148.—Holmia kjerulfi, Linnars. ×

1. Lower Cambrian. (After Holm.)
Fig. 149.—Clenelloides armatus, Peach.
Lower Cambrian, × 3. (After Peach.)
Fig. 150.—A, Harpes ungula, Sternb., Ordovician. B,
Ellipsocephalus hoffi, Scloth., Cambrian. C, Olenus truncatus,
Brünn., Cambrian. (After Angelin.) D, Remopleurides radians,
Barr., Ordovician. E, Conocoryphe sulzeri, Barr., Cambrian. F,
Illaenus dalmanni, Volb., Ordovician. G, Proëtus bohemicus,
Corda, Silurian, × 1½. H, Aeglina prisca, Barr., Ordovician, × 3.
I, Phacops sternbergi, Barr., Devonian. (A, D, E, G, H, I, after
Barrande; B, F, from Zittel; natural size except G, H.)
 Fam. 9. Asaphidae (Fig. 150, F).—The body is oval and
commonly rather large. The cephalic shield is large, with its glabella
often indistinctly limited and the glabella-furrows often obscure. The
facial suture starts from the posterior margin and usually cuts the
anterior margin, but is sometimes continued in front of the glabella.
The relative size of the fixed and free cheeks varies greatly. The eyes
are of variable size. The thorax consists of eight or ten (sometimes
fewer) segments; the pleurae are generally grooved, but sometimes
plane. The pygidium is large, often being similar in form and size to
the head; it consists of numerous segments which, however, may be
indistinctly shown; the axis in some forms is obsolete. Upper
Cambrian (Tremadoc) to Silurian; common in the Ordovician.
Genera: Asaphus (sub-genera, Megalaspis, Asaphellus,
Symphysurus, etc.), Ogygia, Barrandia, Niobe, Nileus, Illaenus,
Bumastus, Stygina. Aeglina (Fig. 150, H) is usually placed in this
family, but its systematic position is doubtful.
Fam. 10. Bronteidae.—The general form is similar to that of the
Asaphidae. The glabella broadens rapidly in front, and is marked
with furrows on each side, which are usually short, and may be
indistinct. The facial suture passes from the posterior margin to the
crescentic eye which is situated rather near the posterior border, and
from thence to the anterior margin. There are ten thoracic segments
with ridged pleurae. The pygidium is longer than the head, and has a
very short axis, from which the furrows on the pleural part radiate.
Ordovician to Devonian. Genus: Bronteus.
Fam. 11. Phacopidae (Figs. 138; 150, I; 151, C).—The head and
pygidium are of about the same size. The glabella is distinctly
limited, and wider in front than behind, with a neck-furrow and
three other furrows, of which some of the anterior may be indistinct
or obsolete. The eyes are schizochroal and usually large. The facial
suture begins at the lateral margin and unites with the suture of the
other side in front of the glabella. There are eleven thoracic segments
with grooved pleurae. The pygidium is usually large, with a distinct
axis and many segments. Ordovician to Devonian. Genera: Phacops,
Trimerocephalus, Acaste, Pterygometopus, Chasmops, Dalmanites,
Cryphaeus.
 Fig. 151.—A, Phillipsia gemmulifera, Phill., Carboniferous. B,
Arethusina konincki, Barr., Ordovician. C, Dalmanites limulurus,
Green, Silurian. (After Hall.) D, Cheirurus insignis, Beyr.,
Silurian. E, Deiphon forbesi, Barr., Silurian. F, Acidaspis
dufrenoyi, Barr., Silurian. (A, B, from Zittel; D, E, F, after
Barrande; natural size.)

Fam. 12. Cheiruridae (Fig. 151, D, E).—The glabella is convex or

inflated, and distinctly defined. The facial suture passes from the
lateral to the front margin. The free cheeks are small, and the eyes
usually rather small. There are from nine to eighteen (usually eleven)
thoracic segments; the pleurae have ridges or grooves and free ends.
The pygidium is small, consisting of from three to five segments
often produced into spines. Upper Cambrian to Devonian. Genera:
Cheirurus, Deiphon, Placoparia, Sphaerexochus, Amphion,
Staurocephalus.
Fam. 13. Proëtidae (Figs. 150, G; 151, A, B).—The body is rather
small, and the head forms about a third of its entire length. The
glabella is sharply defined, and its furrows are sometimes indistinct;
the posterior furrow curves backward to the neck-furrow, thus
limiting a basal lobe on each side of the glabella. The eyes are often
large (Fig. 150, G); but in Arethusina (Fig. 151, B), in which an eye-
line is present, they are small. The facial sutures pass from the
posterior to the anterior margin. The free cheeks are large. There are
from eight to twenty-two thoracic segments with grooved pleurae.
The pygidium is usually formed of numerous segments, and its
margin is usually entire. Ordovician to Permian. Genera: Proëtus,
Arethusina, Cyphaspis, Phillipsia, Griffithides, Brachymetopus,
Dechenella.[203]
Fam. 14. Encrinuridae.—The cephalic shield is ornamented
with tubercles. The free cheeks are narrow, and the eyes very small.
The facial suture extends from the lateral margin (or from the genal
angle) to the anterior margin. There are from ten to twelve thoracic
segments with ridged pleurae. On the axis of the pygidium numerous
segments are seen, but usually fewer are indicated on the lateral
parts. Ordovician and Silurian. Genera: Encrinurus, Cybele,
Dindymene.
Fam. 15. Acidaspidae (Fig. 151, F).—The cephalic shield is
broad, with a spinose margin, genal spines, and sometimes spines on
the neck-ring. The glabella has a longitudinal furrow on each side,
due to the backward bending of the lateral furrows. The facial suture
passes from the posterior border (near the genal angle) to the
anterior border. The free cheeks are large; the eyes small. There are
from eight to ten thoracic segments with ridged pleurae, which are
produced into long backwardly directed spines. The pygidium is
short, and is formed of two or three segments with long spines at the
margin. Ordovician to Devonian. Genus: Acidaspis.