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Sperm Morphology of Domestic Animals
Sperm Morphology of Domestic Animals

J.H. Koziol, DVM, MS, DACT

Associate Professor of Food Animal Medicine and Surgery
Texas Tech University School of Veterinary Medicine
Amarillo, TX, USA

C.L. Armstrong, DVM, MS, DACT

Associate Veterinarian
Elgin Veterinary Hospital
Elgin, TX, USA
This edition first published 2022
© 2022 John Wiley & Sons, Inc.

All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted,
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The right of J.H. Koziol and C.L. Armstrong to be identified as the authors of this work has been asserted in
accordance with law.

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John Wiley & Sons, Inc., 111 River Street, Hoboken, NJ 07030, USA

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Library of Congress Cataloging-­in-­Publication Data applied for

ISBN: 9781119769767 (hardback)

Cover Design: Wiley

Cover Image: Courtesy of J.H. Koziol, Westend61/Getty Images

Set in 9.5/12.5pt STIXTwoText by Straive, Pondicherry, India

10 9 8 7 6 5 4 3 2 1
For RLC and DFW who taught us to never be satisfied and complacent with the status quo
 vii

Contents

Preface ix
About the Companion Website x

Introduction 1

Section I Head Abnormalities 11

1 Pyriform and Tapered Heads 13

2 Nuclear Vacuolation, Including the Diadem Defect 20

3 Macrocephalic and Microcephalic Sperm 29

4 Rolled Head–Nuclear Crest–Giant Head Syndrome 32

5 Abnormal DNA Condensation 34

6 Acrosome Abnormalities 36

7 Normal Detached Heads and Free Abnormal Heads 44

8 Decapitated Sperm Defect 47

Section II Midpiece Abnormalities 49

9 Proximal Cytoplasmic Droplet 51

10 Pseudodroplet 58

11 Mitochondrial Sheath Defects 59

12 Corkscrew Sperm Defect 65

viii Contents

13 Dag Defect 68

14 Distal Midpiece Reflex 72

15 Bowed Midpieces 78

16 Distal Droplets 79

17 Abaxial Midpieces 82

Section III Tail (Principal Piece) Abnormalities 85

18 Tail Stump Defect 87

19 Coiled Principal Pieces 90

20 Double Forms and Accessory Tails 93

21 Bent Principal Pieces 97

22 Short Tail Defect 99

Section IV Aberrations of Stains and Other Cells in the Ejaculate 101

23 Aberrations Due to Staining 103

24 Aberrations Due to Cold Shock 106

25 Round Cells in the Ejaculate 107

26 Teratoids 110

27 Medussa “Cells” 112

28 Aberrations of Semen Quality 113

References 116
Index 125
 ix

Preface

Sperm Morphology of Domestic Animals has been written to further the understanding of morphologic
assessment in the dog, stallion, bull, buck, and ram. This text is directed toward veterinary students
and practitioners. Morphologic evaluation of sperm is a confusing part of the fertility examination.
Ample images and figures have been added to provide real-­time support during semen evaluation.
The authors sincerely hope the added imagery will help provide clarity to a complex and often
confusing portion of fertility assessment. Morphologic evaluation of sperm is an ever-­evolving area
of theriogenology, and this text has been written with the latest information from the current
literature.
The authors would like to thank their mentors and other leaders in the field of theriogenology
that inspire excellence in all that they set out to accomplish. This text would not have been possible
without the generous support from Wiley.
x

About the Companion Website

This book is accompanied by a companion website:

www.wiley.com/go/koziol/sperm

The website includes:

●● Teaching PowerPoints
 1

Introduction

Sperm evaluation provides a noninvasive method to evaluate testicular and epididymal function,
providing information similar to that gained by a testicular biopsy. An abnormal spermiogram
with supporting evidence from the history and physical exam can give insights into reasons for
abnormal testicular function, and consequently allow formation of a prognosis for recovery or
potential treatment. When an abnormal spermiogram is found, the types and number of abnor-
malities combined with history regarding environment, nutrition, and health status can be used to
compile a reason for spermiogram disturbances noted. The veterinarian can then use that informa-
tion to make a diagnosis and prognosis for recovery.
An awareness of the mechanism of sperm transport in the female reproductive tract aids in
understanding the relationship between sperm quality and fertility and consequently the level of
defects that can be tolerated and still maintain reasonable pregnancy rates. The cervix, uterus, and
uterotubal junction all reduce the number of abnormal sperm that can reach the site of fertiliza-
tion. The cervix filters sperm with tail defects, while severely deformed heads are filtered at the
level of the cervix, uterus, and uterine tubes. However, the filter system is not perfect and some
sperm abnormalities may still reach the oocyte and induce fertilization [1, 2].
Three main criteria should be taken into account when evaluating abnormal sperm morphology:
(i) abnormalities of the nucleus that allow ovum penetration and zona reaction but not fertiliza-
tion or embryonic development cannot be tolerated at levels >15–20% without subsequent decrease
in pregnancy rate; (ii) abnormalities of the acrosome and sperm tail do not interfere with the abil-
ity of normal sperm to fertilize ova and therefore can be tolerated at levels up to 25%; (iii) a least
70–75% of spermatozoa should be normal [3]. The types and percentages of defects should be
taken into consideration in both natural and artificial mating systems.

I.1 ­Spermatogenesis

A thorough understanding of normal and abnormal spermatogenesis facilitates understanding and

interpretation of a spermiogram (description of sperm morphology noted during evaluation)
(Figures I.1–I.4). The testis is largely composed of seminiferous tubules and interstitial tissue. The
latter is located between the seminiferous tubules and consists primarily of Leydig cells, which pro-
duce and secrete steroid hormones, as well as vascular and lymph vessels that supply the testicular
parenchyma. Seminiferous tubules arise from primary sex cords and contain germinal cells, as well
as Sertoli cells, which support and nurture production of sperm. Sertoli cells form tight junctions,

Sperm Morphology of Domestic Animals, First Edition. J.H. Koziol and C.L. Armstrong.
© 2022 John Wiley & Sons, Inc. Published 2022 by John Wiley & Sons, Inc.
Companion website: www.wiley.com/go/koziol/sperm
 2 Sperm Morphology of Domestic Animals

Figure I.1 Morphologically normal sperm from a bull (eosin–nigrosin, 1000×).

creating the most important portion of the blood–testis barrier [4]. Spermatogenesis occurs primar-
ily in the tubulus contortus section of the seminiferous tubules. These tubules are surrounded by
contractile peritubular cells that promote flow out of the tubulus contortus into the rectus (straight
portion) of the seminiferous tubule. Both ends of each tubule are connected with the rete testis.
Spermatogenesis can be divided into three phases. The first phase, proliferation, consists of
six mitotic divisions of spermatogonia, increasing the number of A-­spermatogonia then
B-­spermatogonia. An important part of this phase is stem cell renewal. Loss of intercellular bridges
allow some spermatogonia to revert to stem cells. B-­spermatogonia then undergo several mitotic
divisions, with the last division resulting in primary spermatocytes. The meiotic phase begins with
diploid primary spermatocytes. During meiosis I, genetic diversity is ensured by DNA replication
and crossing over during production of secondary spermatocytes, and as a result, from a genetic
perspective, no two sperm are identical. The last phase, known as the differentiation phase or sper-
miogenesis, is differentiation of a spherical undifferentiated spermatid into a fully differentiated
sperm with a head, a flagellum which includes the midpiece, and the principal piece. The length
of spermiogenesis differs between species. An example is that it takes the last 18 days for the sper-
miogenesis stage of spermatogenesis to be completed in the bull.
Correct clinical interpretation of spermiograms also requires understanding of the specific tim-
ing of spermatogenesis in the species being evaluated. An understanding of the cycle of the semi-
niferous epithelium aids in this mission. The cycle of seminiferous epithelium is the progression
through a complete series of stages at one location along a seminiferous tubule. On cross-­sectional
evaluation of the seminiferous tubule, four to five concentric layers of germ cells are present with
each layer representing a generation. Each cross-­section along the length of the seminiferous
tubule will have a distinct appearance. Each cross section with its four to five generations of germ
cells represents a stage of the seminiferous epithelium cycle. For example, a cross section in stage
1 will have a base layer of A-­spermatogonium followed by primary spermatocytes, and another
layer of primary spermatocytes, followed by a layer of immature spermatids. A stage 8 cross sec-
tion will begin with a layer of A-­spermatogonium, followed by B-­spermatogonium, primary

0005236578.INDD 2 12-15-2021 16:44:14

 Introduction 3

Figure I.2 Morphologically normal sperm from a bull (phase contrast wet mount, 1000×).

spermatocytes, immature spermatids, and mature spermatids that are ready to be released into the
lumen. Along the length of any given seminiferous tubule, there are many zones or cross sections
in different stages with only a few zones at an appropriate stage to release mature spermatid into
the lumen of the seminiferous tubule, which will then travel through the rete testis to be trans-
ported to the head of the epididymis. These different zones allow for the creation of a spermato-
genic wave or constant release of sperm along the length of each tubule. With zones in multiple

Figure I.3 Morphologically normal sperm from a buck (eosin–nigrosin, 1000×).

 4 Sperm Morphology of Domestic Animals

Figure I.4 Morphologically normal sperm from a stallion (eosin–nigrosin, 1000×).

stages along the length of the tubule, there is a constant flow of spermiation, which allows for a
constant flow of sperm to the epididymis.
Normal testicular function is dependent on appropriate endocrine, autocrine, and paracrine con-
trol. In the male, gonadotropin releasing hormone (GnRH) is secreted from the hypothalamus in a
pulsatile nature resulting in waves of luteinizing hormone (LH) and follicle stimulating hormone
(FSH) release. LH acts on the Leydig cells within the testes, which are responsible for synthesizing
progesterone, the majority of which is converted to testosterone. This pulsatile nature of LH is neces-
sary for normal testicular function as sustained LH can lead to a refractory nature of the Leydig cells
secondary to a downregulation in the number of receptors. This culminates in reduced secretion of
testosterone from the Leydig cells. High concentrations of testosterone must be maintained for normal
spermatogenesis to occur. The pulsatile release of LH and consequently testosterone prevents negative
inhibition of FSH, which is needed for maintenance of Sertoli cells. Sertoli cells convert testosterone
to dihydrotestosterone and estradiol along with secreting androgen binding protein (ABP).
Approximately 80% of the secreted ABP goes into the lumen of the seminiferous tubule and travels to
the epididymis, while the remainder is secreted into the interstitial compartment and is absorbed into
the systemic circulation. ABP binds testosterone and is responsible for not only maintaining a high
concentration of testosterone within testes but also carries testosterone to the epididymis.
The most common causes of abnormal spermatogenesis in males include abnormal testicular
thermoregulation; hormonal imbalances, particularly those associated with stress; and effect(s)
of toxins or expression of deleterious genes [5]. Stress typically elevates systemic cortisol concen-
trations, profoundly decreasing release of LH and testosterone [6–8]. Stress has many origins,
including environment, illness, or injury, causing changes in the spermiogram similar to those
induced by disruption of thermoregulation. The primary spermatocyte is extremely sensitive to
alterations in the hormonal milieu secondary to stress or illness. For example, in cases of testicu-
lar degeneration, the changes appear first in the cytoplasm, centrosomes, and spindles at the level
of the primary spermatocyte, which predispose to disturbances in the developing spermatid [3, 9].

0005236578.INDD 4 12-15-2021 16:44:17

 Introduction 5

I.2 ­Length of Spermatogenesis in Domestic Species

Stage Bull [10] Ram [10] Boar [10] Stallion [10] Dog [11]

I 4.2 2.2 1.1 2.0 3.0

II 1.2 1.1 1.4 1.8 1.7
III 2.7 1.9 0.4 0.4 0.4
IV 1.7 1.1 1.2 1.9 1.6
V 0.2 0.4 0.8 0.9 1.1
VI 0.8 1.3 1.6 1.7 2.1
VII 1.1 1.1 1.0 1.6 1.8
VIII 1.6 1.0 0.8 1.9 1.9
Length of one cycle 13.5 10.4 8.3 12.2 13.6
Total days for 61 47 39 57 61
spermatogenesis

I.3 ­Maturation of Spermatozoa in the Epididymis

From the rete testis, sperm travel to the efferent duct and into the head of the epididymis. Grossly,
the epididymis can be divided into three segments: the head (caput), body (corpus), and tail
(cauda). The epididymis is responsible for transport, concentration, storage, and maturation
of sperm.
During epididymal transport, maturation of sperm continues with many biochemical and mor-
phological modifications occurring. All sperm that enter the head of the epididymis do so with a
proximal cytoplasmic droplet; as they move through the epididymis, the droplet normally migrates
down the midpiece and is present at the distal end of the midpiece when sperm arrive in the tail of
the epididymis.
Along the entire length of epididymis, the lumen is bordered by epithelium that synthesizes and
secretes proteins under androgen stimulation [12]. Intraluminal fluids released sequentially along
the epididymal lumen are key for maturation of sperm [13]. As the sperm cell travels along the
lumen many biochemical modifications occur. These processes include nucleus chromatin con-
densation, changes in plasma membrane composition and relocation of surface antigens, and the
ability to respond to hypo-­osmotic stress [14]. Circulating and intraluminal androgens are critical
for epididymal gene expression and secretion. Androgens in the epididymis are synthesized by
enzymatic reduction of testicular testosterone. Estrogen produced by the testicular Leydig cells
also plays a role in the epididymis particularly in the role of water reduction in the proximal regions
of the epididymis [14].
Following ejaculation, distal droplets should be shed after mixing with seminal fluid. Epididymal
transport time is species dependent (Table I.1). The tail of the epididymis and ductus deferens
serve as short-­term sperm storage. However, in the absence of ejaculation, senescent sperm should
be continually expelled into the urinary tract, ensuring that viable sperm are always available, even
following long periods of sexual rest [15].
 6 Sperm Morphology of Domestic Animals

I.4 ­Staining Techniques and Evaluation of Morphology

Microscopic evaluation of sperm morphology is a pillar of successful sperm evaluation and breed-
ing soundness examinations of any species. The three basic types of light microscopy are bright-
field, phase contrast, and differential interface contrast (DIC). Most conventional microscopes
have a brightfield function and many also have phase contrast capability. However, DIC is expen-
sive and is generally restricted to use in research or andrology laboratories.
Brightfield microscopy is suitable for examination of morphology utilizing stained semen
smears. Morphology evaluations must always be performed using the oil immersion (100×) objec-
tive lens; with eyepieces of 10–12.5×, the total magnification is 1000–1250× (hereafter referred to
as 1000×). Morphology evaluation at lower magnification is not recommended as it results in fail-
ure to recognize many sperm abnormalities.
Utilization of phase contrast microscopy allows examination of unstained, cover-­slipped, prepa-
rations prepared by mixing semen with warmed 10% neutral buffered formalin on a microscope
slide, as well as slides stained with Feulgen stain [20]. Evaluation of sperm morphology utilizing
phase contrast at 1000× is preferred over the use of stained preparations by some practitioners, as
defects such as nuclear vacuoles, [21] acrosomal defects, and abnormal DNA in the case of Feulgen-­
stained slides are easier to identify. However, these defects can also readily be identified with high-­
quality brightfield microscopes.
Morphologically abnormal sperm have long been associated with male infertility, and assess-
ment of these abnormalities is a fundamental component of analysis of semen quality. [3, 22–24].
Sperm morphology is an excellent predictor of the outcome of natural mating, artificial insemina-
tion, and in vitro fertilization, [3, 25, 26] and there is a relationship between morphologically
abnormal sperm and poor DNA quality [24].

I.5 ­Vital Staining

Live/dead stains, also referred to as supravital stains, such as eosin–nigrosin and eosin–aniline
blue, are commonly used for routine morphological examination of sperm [27]. In a study by
Tanghe et al., eosin–nigrosin staining of sperm was the only bull sperm quality parameter evalu-
ated that demonstrated a significant association with pronucleus formation in in vitro fertiliza-
tion [28]. When utilizing live/dead stains, nigrosin or aniline blue provides a background, whereas
eosin acts as a vital stain. Functional plasma membranes of viable sperm prevent penetration of
eosin into the cell; therefore, viable sperm appear white. In contrast, eosin penetrates damaged cell
membranes, staining injured or nonviable sperm pink.

Table I.1 Mean time of sperm epididymal passage in domestic

Species Average time (days) References

Dog 15 Linde-­Forsberg [16]

Boar 9–14 Franca and Cardoso [17]
Bull 8–11 Barth and Oko [3]
Stallion 8–11 Swierstra [18]
Ram 16.4 Amann [19]

0005236578.INDD 6 12-15-2021 16:44:17

 Introduction 7

Figure I.5 Partially stained sperm with a

pink posterior and a white anterior
(eosin–nigrosin, bull, 1000×).

Partially stained sperm, with a pink posterior portion and a white anterior portion of the sperm
head, are sometimes observed. These sperm have suffered membrane damage, yet the acrosome
remained intact. This staining pattern occurs because the acrosomal membrane is tightly applied
to the nucleus at the equatorial region and penetration of the eosin stain under the acrosome is
initially impeded at the equatorial region [29]. Partially stained sperm are more common when
semen samples are processed under less than ideal conditions, suggesting that sperm membrane
damage or cell death occurred very recently (Figure I.5).
When assessing the proportion of sperm staining alive, it may be relevant to only count sperm
stained completely white. When semen has been handled properly and stained correctly, the per-
centage excluding eosin, and therefore considered “alive” is highly correlated to progressive motil-
ity. This is not to suggest that only the “alive” cells be counted during evaluation of a
sperm morphology slide; 100 cells should be counted regardless of vital status to give an
accurate representation of the current state of the ejaculate. The accuracy of the assess-
ment of marginal bulls can be improved by counting 300 cells.

I.6 ­Preparation of Semen Smears or Coverslip Slides

Slide preparation is crucial for proper evaluation of sperm morphology and requires practice and
attention to detail. Common errors include too many sperm per slide (Figure I.6); slides stained too
dark or too light; and slow drying resulting in cracks and stain artifacts. There are multiple ways to
create stained slides. Some may prefer to streak out slides using another glass slide in a manner simi-
lar to producing a blood smear while others prefer to use a wood applicator stick to roll the sample
out (Figures I.7–I.9). Some practitioners believe that the wood applicator stick results in less mechan-
ical damage to the sperm (detached heads, damaged acrosomes). However, no studies have proven
this clinical suspicion. Slides can be viewed anywhere along the length but usually half way down the
slide is a good place to start as the cells are usually evenly distributed with good background stain.
Wet mounts can be made for evaluation of sperm morphology by phase-­contrast or DIC micros-
copy at 1000×. Freshly ejaculated semen must be diluted and motility stopped to allow for accurate
evaluate. This can be accomplished with the addition of 0.2% glutaraldehyde in phosphate buffered
saline (PBS), buffered formal saline, or with neutral buffered formalin. To prepare a wet mount
slide, a 2–3 mm drop of diluted semen is placed on a warm clean microscope slide and a coverslip is
placed over top. Allow the drop to evenly distribute under the coverslip before placing a drop of oil
 8 Sperm Morphology of Domestic Animals

Figure I.6 Poorly prepared slide with too many sperm per field, which impeded the evaluators ability to
evaluate each individual sperm (eosin–nigrosin, 500×).

Figure I.7 A small line of stain is placed

along the slide followed by a small drop
of semen.

Figure I.8 Using a second slide, one can

start in the stain backing up to pick up a
small line of semen and then proceed to
streak out the slide. There are many
different methods than can be used for
streaking out slides.

0005236578.INDD 8 12-15-2021 16:44:22

 Introduction 9

Figure I.9 Slides should neither be too

dark or too light, and when viewed under
the microscope, the cells should be spaced
in such a manner that they can each be
individually examined.

on the coverslip for examination at 1000×. The preparation should be very thin to avoid sperm lay-
ing on their edges rather than flat. When using glutaraldehyde or neutral-­buffered formalin, clump-
ing of cells may occur. Wet mount slides can be viewed at any location along the coverslip with the
exception of near the edges, as this is where a higher number of dead cells tend to accumulate [3].

I.7 ­Differential Counting of Sperm Morphology

Differential counts of sperm morphology must be performed using 1000× magnification (oil immersion)
as lower magnifications result in failure to recognize some defects. The goal for examining sperm
morphology is to determine the percentage and types of sperm abnormalities present in a sample and
thereby construct a spermiogram. The spermiogram may be used to interpret potential fertility of
semen at the time of sampling, as well as determine potential causes of an abnormal spermiogram.
Knowledge of probable cause of abnormalities may allow the practitioner to develop a prognosis for
recovery, which will aid the client in making an informed decision about the male in question [29].
Determination of a spermiogram is facilitated by the use of a differential cell counter. Mechanical
cell counters often allow one to press two or more keys simultaneously to record more than one
defect within the same sperm, while counting the individual sperm only once. With this system, a
more accurate spermiogram may be produced.
When sperm morphology is good, differential counting of 100 sperm is adequate to determine a
spermiogram. However, when the percentage of abnormal sperm present is high, counting at least
300 sperm may be necessary to produce an accurate spermiogram.
 11

Section I

Head Abnormalities
 13

Pyriform and Tapered Heads

The prevalence of pyriform head defects are relatively high and are either the most or second most
prevalent defect in an ejaculate [3, 30–34]. The classical pyriform sperm has a pear-­shaped head, a
normal acrosome, and a narrow post-­acrosomal region [3, 35–37]. Sperm with irregularity in head
shape and size generally have good motility [3].
Several variants of pyriform head defects have been reported, ranging from slightly affected to
severely tapered through the acrosome and post-­acrosomal regions. The main initiating factor for
this deformity appears to be adverse environmental conditions impacting male health and fit-
ness [3, 38]. Pyriform sperm have been noted in ejaculates three to four weeks after initiation of
dexamethasone injections to simulate stress, following scrotal insulation, or exposure to high
ambient temperatures in the case of rams [3, 5, 38]. Pyriform or tapered heads have also been noted
in those suffering from obesity resulting in scrotal or inguinal deposition of fat, frostbite or derma-
titis of the scrotum, varicoceles, or other injuries that impede proper testicular thermoregula-
tion [3]. Disruptions in endocrine hormone balance occurring either systemically or locally may
also be implicated. Local disruption is caused by abnormalities in testicular thermoregulation
while systemic disruption descends from unfavorable conditions or events that cause stress to the
animal. These stressful events can include chronic pain, joint pain, conditions affecting the hooves
including lameness or laminitis, extremes in weather, housing or social stress, improper nutrition,
or travel [3].
Differences in the severity of response between males with similar treatments suggest that some
males may be genetically predisposed to develop pyriform heads in response to adverse environ-
mental influences [3, 5, 29]. In most instances, there will be many other sperm abnormalities pre-
sent in the ejaculate, indicating a disturbance in spermatogenesis has occurred. However, after
removal of stress factors, males often return to normal sperm production [3].
Small numbers of pyriform or tapered defects can be found in the semen of most bulls, even
in bulls of good fertility [3]. In a study of bulls used for natural service and artificial insemina-
tion, percentage of bulls producing affected sperm were 9 and 16%, respectively. The percentage
of pyriform sperm in an ejaculate from affected individual varies from 10 to >50% [3]. When
pyriform defects are in high numbers, their impact on fertility has been documented in both
in vivo and in vitro studies. Thundathil et al. [39] reported that in vitro fertilization rates were
lower for semen containing high numbers of pyriform heads than for ejaculates containing low
numbers, 68.5 versus 84.4%, respectively. Sixty-­day pregnancy rates in estrus-­synchronized arti-
ficially inseminated heifers using semen from these same bulls were 37 and 61%, respectively, for
bulls with pyriform sperm and the control bull. In the same study, rates of embryo/fetal loss

Sperm Morphology of Domestic Animals, First Edition. J.H. Koziol and C.L. Armstrong.
© 2022 John Wiley & Sons, Inc. Published 2022 by John Wiley & Sons, Inc.
Companion website: www.wiley.com/go/koziol/sperm
14 Sperm Morphology of Domestic Animals

between rates of embryo/fetal loss between days 22 and 60 of pregnancy were 23 and 8% for the
pyriform head group and the control group, respectively [39].
Pyriform sperm disrupt various stages of the fertilization process including: sperm trans-
port, [40–42] oocyte binding and zona penetration, [43] along with postfertilization events [39].
Furthermore, the prognosis for recovery is highly variable among animals. Mature bulls with
high levels (25–75%) of pyriform sperm have a poor prognosis for recovery when there is no

Figure 1.1 Pyriform head in a bull (eosin–nigrosin, 1000×).

Figure 1.2 Pyriform head in a bull as indicated by arrow (eosin–nigrosin, 1000×).

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Maotente, 103
Fueiros, 119
Cunheiras, 121
Espelhos, 125
Revindo, 126, 128, 287
A pe quedo, 132
Preguntas d’estrada, 135
Salsa de Palmela, 137
Era a rodo, 137
Reguengnos, 149
Boinhos, 150
Bregios, 156
Jogo de malham, 159, 167
Desviados, 165
Mosseos, 167
Sonaves, 181
Cofos, 185
Pelote de usteda, 187
Ceroules, 188
Cervilhas, 188
Bouçaes, 200
Vivos, 216
Enxarafas, 232
Comeeira, 244, 260, 337
En cocras, 247
Alambeis, 266
Releixo, 287
Hopa, 296
Humiziados, 297
Menuta, 314
Escodilhinas, 332
Moucho, 333
Sem tresposta, 336
Cerolho, 340
Revezado, 343
De mao em mao, 365
Impidosa, 384
T. RICHARDS, PRINTER, 37, GREAT QUEEN STREET.
 FOOTNOTES:
[1] See the St. James’s Gazette. Criticism of Mr. Justin
McCarthy’s language, October 29th, published since the above
was written.
[2] Sic.
[3] Or “of the Vision”, according to some.
[4] Caspar Correa says it was a breviary on parchment, and
describes a picture contained in it on paper which had come from
Jerusalem or Rome.
[5] Gaspar Correa says that at this leave-taking the
Portuguese sounded their trumpets and drums and the fleet
saluted, and a ball from a camel gun made three rebounds
amongst the men of the Barnegaes, without hitting anyone. The
Governor sent to ask pardon of him, saying it was a mistake of
the bombardier. The Barnegaes answered that no one was safe
unless God pleased, and that the ball had done no one any harm.
[6] Gaspar Correa says of the Embassy that Don Rodrigo de
Lima was a man of good presence, and a fit man for such a
service; that Jorge d’Abreu, the second person of the Embassy,
was a very well dressed gentleman; that Francisco Alvarez was a
very prudent man, and well informed in all matters of the altar and
divine services; that Estevan Palharte was a good fencer; the
three Portuguese who accompanied Mattheus were Miguel
Fernandez, Diogo Tatys, and João d’Alvarenga, all men skilful in
manual arts, and who could sing at mass, for which the priest
carried rich ornaments and all that was necessary, and irons for
making wafers.
[7] Anta.
[8] “Matos idiabrados”, not in Bluteau.
[9] Meijoada, not in Bluteau.
[10] Oporto.
[11] Alosna.
[12] Como em chacola, dancing and singing, i.e., not
reverentially.
 [13] Como quem anda per mãos diante da imagem. According
to the Spanish translation of Miguel de Salves. Saragossa, 1561.
[14] Oussia.
[15] Under a canopy.
[16] Symam.
[17] By threes.
[18] Mansfield Parkins calls it teff, the species of corn most
esteemed in this country.
[19] Zamboa, not referred to in Selves’ translation.
[20] Should be teskar.
[21] Castar should be Teskar; Mansfield Parkyns calls it a sort
of funereal feast, where charities are bestowed on the poor and
the priests.
[22] Couto, an asylum, an enclosure; celeiro, a granary.
[23] Zagonaes. Ethiopian corruption of diakonos.
[24] Dibarua, hillock seen from afar.
[25] Bait en-negus.
[26] Azorague, a kurbach, or whip of hide, cravache.
[27] A Portuguese measure equal to about three pints.
[28] Padrinho.
[29] Shoa.
[30] Rumană Wark, the golden pomegranate.
[31] This opinion of the Abyssinians appears to have
anticipated the Sorbonne judgment of 1733, given in the 20th
chapter of Tristram Shandy.
[32] Crisma.
[33] Envorilhados for enrodilhados.
[34] Adens and marrecas.
[35] No las assombram a terra: perhaps this means with a
paper kite. This passage is omitted in Selve’s translation.
[36] Cevado, literally, allured.
[37] The favourite or preferred, Bitwaddad, because he was
liked.
 [38] Ar-ras.
[39] Mount Sinai.
[40] Foy desordenado e feito leigo; not a correct expression.
[41] A gold coin in some parts of Africa.
[42] Bechugarias.
[43] Zafoens or Safoens.
[44] Tufas.
[45] Nubians.
[46] Dafilla.
[47] Igzio maranna Kristos, Amharic.
[48] The Latin text of a similar excommunication of noxious
animals was given in the Pall Mall Gazette, or Times, in 1879.
[49] Kurbayra.
[50] Tigri mahom, should be Tigri măkuănăm.
[51] “Where”, should be “here”; this river, the Mareb, not the
Nile, separates the country of the Barnagais from that of Tigré.
[52] This and other passages show that some of the churches
in Abyssinia, which are sometimes called Portuguese churches
by travellers, are anterior to the Portuguese.
[53] Sic.
[54] This passage appears to have been inserted here by
some error of the printer, and to belong to some other chapter.
[55] Pontões.
[56] Sobarbava.
[57] Pliny says Candace was the name of many Queens, as
Pharaoh was of the Kings of Egypt.
[58] Mansfield Parkyns calls it a sycamore tree, and says it is
remarkable for the extraordinary circumference of its trunk and
the great spread of its branches, which cast their shade over such
a space of ground as would be sufficient for the camp of the
largest caravan.
[59] Mansfield Parkyns thus describes it: “The principal obelisk
is carved on the south side, as if to represent a door, windows,
cornices, etc.”
 [60] Mancal, an unknown game, antiquated in the reign of Don
Manuel; the translation of Selves calls it a juego de herradura.
“Baton ferré des deux bouts”.—Roquette’s Dict.
[61] Arouca, a town in the bishopric of Lamego.
[62] Nebrete, i.e., Nabrĭd, imposed by hand, the title of the
Governor of Axum.
[63] Maabar or Mahabar, a club, gathering.
[64] Feitos igoaes; the Selves translation has, las camas
hechas, y por cubrir.
[65] Abacinete, i.e., Amba Sanete, name of a hill.
[66] Estefarruz is probably a misprint of Alvarez for Estefanuz,
or Stephen.
[67] i.e., Fillets.
[68] Maluk.
[69] Or Kyriakos.
[70] Gada is an expedition sent from the highlands to collect
the salt, Băal means master or chief; Băalgada is a title given to
the chiefs of those expeditions, and is a title still used in Ethiopia,
like Duke or Count.
[71] Estrevimento, antiquated.
[72] The drachm of gold; that is, the salt increases in value.
[73] Dagusa (Eleusine tocusso), used for bread, but more for
brewing beer. Mansfield Parkyns says (vol. i, p. 265): “very little
‘teff’ is grown in this neighbourhood, but chiefly millet (mashéla)
and dagousha”.
[74] Valuras.
[75] Maotente.
[76] Igzio maranna kristos.
[77] Nug (Guizotia oleifera), Pampilhos, a plant like olho de
boy.
[78] Apodavam.
[79] Cătăma, a camp.
[80] Or Caravan of merchants.
 [81] Conquest here means maintaining the struggle, or the
duty of conquering. Whilst the Moors were still unconquered in
the south of Spain, certain parts of their territory were said to
belong to the Conquest of Castile, others to the Conquest of
Aragon.
[82] Poçoncha.
[83] Twenty-six almudes make a pipe.
[84] Both these towns of Corcora are in the modern maps.
[85] Por amor de unas sierras.
[86] Gabya.
[87] Esparto.
[88] Yĭkun amlak, may he be Lord.
[89] This passage is not very clear. Selves translates: “the
church lies to the east, and the Epistle is said at the west”.
[90] Fueiros, projections over the sides of a cart to enable it to
carry more load, side rails, wings, thripples, shelvings.
[91] Terza, nine o’clock.
[92] Cunheiras, not in the dictionaries, possibly for wedge
marks.
[93] Palmo, measure of four inches.
[94] Corucheo; Selves calls it coroza.
[95] Espelhos.
[96] Frecheiros.
[97] Lunas.
[98] Of the wall of inclosure.
[99] Revindas, Revindo, a, not in any dictionary except the
Diccionario Portatil of Santa Rosa de Viterbo, Coimbra, 1825,
which says: “Arco, on abobeda de meio circulo perfeito que dizem
de meia volta em berço”.
[100] Revinda.
[101] Curral.
[102] A pe quedo.
 [103] Preguntas d’estrada, wayside questions. The question
about the length of time the Holy Family passed in Egypt was not,
however, a simple or purposeless question, as the Abyssinians
assert that during that time they visited Abyssinia.
[104] Salsa de Palmela.
[105] Era a rodo.
[106] Church of the Trinity.
[107] With his knapsack in his hand. Con la barjuleta en la
mano, Selves.
[108] Furou na silveira; Rompiendo por una enramada,
Selves.
[109] In the preceding chapter it was said to be two days’
journey in circumference.
[110] This is the worst trait of Abyssinian character to be found
in any of the accounts of that country; and instances of similar
conduct are only to be found in Russia.
[111] See Major Harris for an account of the imitation by the
King of Shoa of this custom of shutting up the King’s relations.
[112] Esparavel, a parasol or umbrella, apparently a mark of
rank.
[113] Morgados ou coutos.
[114] Reguenguos, private estates of the King.
[115] De ab inicio.
[116] Amhara.
[117] See Cap. 57, p. 139.
[118] Cabaça.
[119] Boinhos.
[120] Gumari, a hippopotamus.
[121] Lixa squatina.
[122] Makana Selase, place of the Trinity.
[123] Bregĭos for brejos.
[124]? West.
[125] Mareados.
 [126] Valuras.
[127] Piçarra.
[128] Joguos de malham. Selves translates: “juegos de
herraduras.”
[129] Espinguam. Selves calls it cuchilla.
[130] Meijoada.
[131] Moyo, modius, measure of sixty alqueires, or bushels.
[132] Probably Debra Libanos.
[133] Maioral.
[134] Ichage is the chief monk of Ethiopia.
[135] Misprinted testar.
[136] Găbăz is a prior in Aksum. There is a Găbăz in every
church, but there the title means the chief of the vestry.
[137] Alqueire.
[138] Cingida.
[139] Hawaryat means apostles, and is thus used in the name
of a church.
[140] Desciados.
[141] Franks; who these were is explained in Cap. 72.
[142] Malham.
[143] Mosseos.
[144] Not translated by Selves.
[145] Bitwaddad, because he was liked, a favourite.
[146] Cacha.
[147] De Xio, a Greek.
[148] Copts. They still call white people Copts; see M.
Parkyns.
[149] Encoiradas.
[150] Selves translates, “when he lowers it”.
[151] Sonaves. Selves has traves.
[152] Băzzăto means carded cotton.
 [153] Cofos.
[154] Cruzados.
[155] Manicordio, also written monocordio. See Grove’s
Dictionary of Music for a note on the Monochord.
[156] Pelote de usteda; not mentioned by Selves: a jacket or
tunic without sleeves.
[157] Ceroules. Selves has çaraguelles.
[158] Cervilhas. Selves has Xervilletas.
[159] Of Nice. Selves has Niceno.
[160] Selves renders this: “the church of Greece had been
founded by the Apostle St. John at Ephesus, and that of
Alexandria by St. Mark the Evangelist.”
[161] The Council of Nice.
[162] This date does not agree with the preceding one, which
would make it Monday.
[163] Alaquequa, Selves translates lazos. “Alaqueca, laqueca,
pierre des Indes qui arrête le flux de sang.”—Roquette’s Dict.
[164] Bouçaes, boçal.
[165] Selves has corrected this to St. Paul.
[166] Maçaã baiones.
[167] Detença.
[168] St. Barlaam, November 19.
[169] 25th November is St. Catharine’s day.
[170] Or fugleman.
[171] Vivos, an ornament for coifs.
[172] Sendeiros Galegos.
[173] Padiola, a hand-barrow.
[174] See Chapter lxv, p. 160.
[175] From this it appears that his leg-bone was not quite
broken.
[176] Sancta sanctorum.
[177] Sombreira, the same word is used for umbrellas or hats.
 [178] Or hollow.
[179] Enxarafas.
[180] A canada equal to three pints.
[181] Adoration of the Magi. Col. Meadows Taylor describes a
similar miracle play represented at Aurungabad by the
Portuguese monks.—Story of my Life, p. 39.
[182] De comeeira, omitted by Selves.
[183] Selves has abridged this chapter considerably.
[184] En cocras, for en cocaras.
[185] Deacons, from diaconos.
[186] This passage is not clear. Selves has, “They put their
hands on the church door as though opening and shutting it.”
[187] From Selves.
[188] Que zagonassem, literally to zagonise.
[189] Of Egypt.
[190] Tero.
[191] Bedem.
[192] Comprida de comieira. See page 244.
[193] Sidy, Sir, Arabic.
[194] Mozo d’espolas.
[195] Alambeis; almayzales, Selves.
[196] The Constant Prince.
[197] Cruzados.
[198] See Gaspar Correa’s account of King John’s scouts in
Vasco da Gama’s Voyages, pp. 8–11.—Hakluyt Society.
[199] The leg-bone, supposed by popular superstition in Spain
to be buried in the great mosque of Cordova.
[200] Jerusalem.
[201] The Portuguese fleet.
[202] Selves translates: “one was wounded in four places by a
gun which must have been loaded with slugs.”
 [203] This passage is very obscure; it either means that the
lords had not seen Jorge d’Abreu and his companions, or that
they did not wish to see them.
[204] The ships of the captain-major.
[205] Milho zaburro.
[206] Guza, probably dagusha.
[207] Agriones.
[208] Revinda.
[209] Releixo.
[210] Pendenças.
[211] Maundy, from the words of Christ, “Hic est mandatum
meum”.
[212] De molde.
[213] In albis.
[214] Pascoela.
[215] Cernache dos alhos, two leagues from Coimbra.
[216] I.e., the termination of the wooden ladder.
[217] Paredinhas.
[218] Seto.
[219] Macho murzelo; mulo morzillo.—Selves.
[220] Hopa, a cloak without sleeves.
[221] Humiziados, Selves translates enfermos, sick people.
Humiziados are persons hiding on account of some crime they
have committed.
[222] Obscure; perhaps the words, “to stay longer”, have been
omitted.
[223] See chapter cix.
[224] Mahfuz.
[225] This figure, xix, is probably a misprint.
[226] Neither Barros nor G. Correa give this date.
[227] Or Seville belonging to Spain, etc., etc.; Selves
translates: “I told him to write Spain under Seville, Galicia under
Corunna, and Portugal under Lisbon.”
 [228] All this passage is omitted by Selves.
[229] Menuta.
[230] Ouquia, a gold coin worth twelve cruzados.
[231] Sultan, apparently the title of a magistrate of Arquigo.
[232] A king’s messenger.
[233] Gulto.
[234] Tribute.
[235] Gaveta.
[236] Or such as is still generally worn by the Aragonese
peasants.
[237] 12,000 men in all.
[238] Gaveta.
[239] See chapter xlviii, p. 110.
[240] Esperavel, should be esparavel.
[241] Corregido; Selves has, la corte ... estaba aderezado.
[242] January 6, Twelfth Day.
[243] The Prester’s ambassador.
[244] Escodilhinas, for escudellinhas.
[245] Mouchas for mochas, without horns, or points, or, in the
case of chairs, may be arms.
[246] Desembargadores.
[247] See chapter cvi, p. 278, where the Abdenago acted in
this way.
[248] Sem tresposta.
[249] Comierias for cumieiras, summit, point of a building. See
pp. 244, 260.
[250] I.e., of the open space going to the market.
[251] Cerolho. This passage is omitted in Selves’ translation.
Cerolho might be a misprint for serodio, late, tardy.
[252] Palmos.
[253] Atabales.
[254] Of Egypt.
 [255] Meijoadas.
[256] Or, alternately, revezadas.
[257] Na falha.
[258] Aduares.
[259] Dankali.
[260] The palace.
[261] Entrudo, Shrove Tuesday.
[262] Or, inhabitants.
[263] Era lhe dado por conto ou camara para hum pequeno
mosteiro; or, “in payment for”, as Selves translates.
[264] Sysones, omitted by Selves.
[265] This man appears to be one of the Prester’s officials.
See before, p. 316.
[266] Golfam.
[267] De mao em mao.
[268] Pardao, Indian coin worth 3 testoons and 3 vintems.
[269] Fortuna.
[270] Gulf of Ormuz.
[271] Selves translates: no hace de noche sino arañar la tierra
y buscar.
[272] The kingdom of Fartaque was on the Arabian coast, east
of Aden. It contained the town of Dhafar.
[273] The 25th of November is St. Catherine’s day, so that
there must be an error as to Saturday or Sunday. Selves avoids
this difficulty.
[274] This superstition still exists among sailors.
[275] Almadia.
[276] Agraço.
[277] Impidosa.
[278] Almotacem-mor.
[279] This would be the gate of the oldest part of the town.
Selves omits all reference to the topography of Coimbra.
 [280] On the coast of India, to the north of Goa.
[281] Milhōes.
[282] Should be seven years.
[283] Akegue, an Emperor.
[284] Pampilhos, also called olho de boy.
[285] Anta is a South American animal.
[286] Probably hyænas.
[287] Adens, marrecas.
[288] Tahu. This is the Tau-cross or T-shaped crutch emblem
of St. Anthony, so called from the name of the letter in the Greek
alphabet.
[289] Tigia.
[290] The Malabar Christians had the same custom. See
Duarte Barbosa.
[291] As mais acavalanas logo.

Transcriber’s Notes:

1. Obvious printers’, punctuation and spelling errors have been corrected

silently.

2. Where hyphenation is in doubt, it has been retained as in the original.

3. Some hyphenated and non-hyphenated versions of the same words have

been retained as in the original.

4. Errata and notes have been silently corrected.

*** END OF THE PROJECT GUTENBERG EBOOK NARRATIVE OF
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