Monotremes diverged from therian mammal ancestors approximately 184 million years ago and have a number of novel reproductive characteristics. One in particular is their penile morphology. There are differences between echidna and platypus phalluses, but both are somewhat similar in structure to the reptilian phallus. The echidna penis consists of 4 rosette glans, each of which contains a termination of the quadrifurcate urethra, but it appears that only 2 of the 4 glans become erect at any one time. Despite this, only a few historical references describe the structure of the echidna penis and none provides an explanation for the mechanisms of unilateral ejaculation. This study confirmed that the echidna penis contains many of the same overall structures and morphology as other mammalian penises and a number of features homologous with reptiles. The corpus cavernosum is well supplied with blood, extends up to the base of the glans penis and is primarily responsible for erection. However, the echidna possesses 2 distinct corpora spongiosa separated by a septum, each of which surround the urethra only distal to the initial urethral bifurcation in the glans penis. Together with the bifurcation of the main penile artery, this provides a mechanism by which blood flow could be directed to only one corpus spongiosum at a time to maintain an open urethra that supplies 2 of the 4 glans to facilitate unilateral ejaculation.

1.
Bassett
EG
.
Observations on the retractor clitoridis and retractor penis muscles of mammals, with special reference to the ewe
.
J Anat
.
1961
;
95
:
61
77
.
2.
Biggers
JD
.
Reproduction in male marsupials
. In:
Rowlands
IW
, editor.
Comparative biology of reproduction in mammals; the proceedings of an international symposium, vol 15
.
London
:
Academic Press
;
1966
. p.
251
80
.
3.
Boas
JEV
.
Zur Morphologie der Begattungsorgane der amnioten Wirbelthiere
.
Morpholog Jahrb
.
1891
;
17
:
271
87
.
4.
Brennan
PL
.
Studying Genital Coevolution to Understand Intromittent Organ Morphology
.
Integr Comp Biol
.
2016
;
56
(
4
):
669
81
.
5.
Cúneo
R
,
Ramezani
J
,
Scasso
R
,
Pol
D
,
Escapa
I
,
Zavettieri
AM
,
.
High-precision U-Pb geochronology and a new chronostratigraphy for the Cañadón Asfalto Basin, Chubut, central Patagonia: implications for terrestial faunal and floral evolution in Jurassic
.
Gondwana Res
.
2013
;
24
(
3-4
):
1267
75
.
6.
Disselhorst
R
.
Die männlichen Geschlechtsorgane der Monotremen und einiger Marsupialen
.
Denkschriften der Medicinisch-Naturwissenschaftlichen Gesellschaft zu Jena
.
1904
;
6_2
:
119
50
.
7.
Dowling
HG
,
Savage
JM
.
A guide to the snake hemipenis: a survey of basic structure and systematic characteristics
.
Zoologica
.
1960
;
45
:
17
28
.
8.
Gerhardt
U
.
Der gegenwärtige Stand der Kenntnisse von den Copulationsorganen der Wirbeltiere, insbesondere der Amnioten
.
Ergebnisse und Fortschritte der Zoologie
.
1909
;
1
:
307
402
.
9.
Goscinski
WJ
,
McIntosh
P
,
Felzmann
U
,
Maksimenko
A
,
Hall
CJ
,
Gureyev
T
,
.
The multi-modal Australian ScienceS Imaging and Visualization Environment (MASSIVE) high performance computing infrastructure: applications in neuroscience and neuroinformatics research
.
Front Neuroinform
.
2014
;
8
:
30
.
10.
Gredler
ML
,
Larkins
CE
,
Leal
F
,
Lewis
AK
,
Herrera
AM
,
Perriton
CL
,
.
Evolution of external genitalia: insights from reptilian development
.
Sex Dev
.
2014
;
8
(
5
):
311
26
.
11.
Gredler
ML
,
Seifert
AW
,
Cohn
MJ
.
Morphogenesis and patterning of the phallus and cloaca in the American alligator, Alligator mississippiensis
.
Sex Dev
.
2015
;
9
(
1
):
53
67
.
12.
Griffiths
M
.
Echidnas
.
Oxford
:
Pergamon Press
;
1968
.
13.
Home
E
.
A description of the anatomy of the Ornithorhynchus paradoxus
.
Phil Trans R Soc Lond
.
1802a
;
92
:
67
84
.
14.
Home
E
.
XI
.
Description of the anatomy of the Ornithorhynchus hystrix
.
Phil Trans R Soc London
.
1802b
;
92
:
348
64
.
15.
Johnston
SD
,
Madden
C
,
Nicolson
V
,
Cowin
G
,
Pyne
M
,
Booth
R
.
Venipuncture in the short-beaked echidna
.
Aust Vet J
.
2006a
;
84
(
1-2
):
66
7
.
16.
Johnston
SD
,
Madden
C
,
Nicolson
V
,
Pyne
M
.
Identifying the sex of short-beaked echidnas
.
Aust Vet J
.
2006b
;
84
(
1-2
):
63
5
.
17.
Johnston
SD
,
Nicolson
V
,
Madden
C
,
Logie
S
,
Pyne
M
,
Roser
A
,
.
Assessment of reproductive status in male echidnas
.
Anim Reprod Sci
.
2007a
;
97
(
1-2
):
114
27
.
18.
Johnston
SD
,
Smith
B
,
Pyne
M
,
Stenzel
D
,
Holt
WV
.
One-sided ejaculation of echidna sperm bundles
.
Am Nat
.
2007b
;
170
(
6
):
E162
4
.
19.
Johnston
S
,
Rumph
C
,
Lucht
M
,
Stenzel
D
,
Nicolson
V
,
Blyde
D
,
.
Studies of Male Reproduction in the Greater Bilby Macrotis lagotis
.
Australian Zoologist
.
2010
;
35
(
2
):
315
30
.
20.
Johnston
SD
,
Lever
J
,
McLeod
R
,
Oishi
M
,
Qualischefski
E
,
Omanga
C
,
.
Semen collection and seminal characteristics of the Autralian saltwater crocodile (Crocodylus porosus)
.
Aquaculture
.
2014
;
422-423
:
25
35
.
21.
Keibel
F
.
Zur Entwickelungsgeschichte des Urogenitalapparates von Echidna aculeata var. typica
.
Denkschriften der Medicinisch-Naturwissenschaftlichen Gesellschaft zu Jena
.
1904
;
6
:
151
206
.
22.
Kelly
DA
.
Intromittant organ morphology and biomechanics: defining the physical challenges of copulation
.
Integr Comp Biol
.
2016
;
56
(
4
):
705
14
.
23.
Larkins
CE
,
Cohn
MJ
.
Phallus development in the turtle Trachemys scripta
.
Sex Dev
.
2015
;
9
(
1
):
34
42
.
24.
Leal
F
,
Cohn
MJ
.
Development of hemipenes in the ball python snake Python regius
.
Sex Dev
.
2014
;
9
(
1
):
6
20
.
25.
Owen R
.
Generative organs of Mammalia
. In:
Owen R
, editor:
On the anatomy of vertebrates, vol 2
.
London
:
Longmans, Green & Co.
;
1868
. p.
641
708
.
26.
Schultz
NG
,
Lough-Stevens
M
,
Abreu
E
,
Orr
T
,
Dean
MD
.
The baculum was gained and lost multiple times during mammalian evolution
.
Integr Comp Biol
.
2016
;
56
(
4
):
644
56
.
27.
Temple-Smith
PD
.
Seasonal breeding biology of the platypus, Ornithorhynchus anatinus (Shaw, 1799), with special reference to the male. PhD thesis
.
Australian National University
;
1973
.
28.
Tyndale-Biscoe
CH
,
Renfree
MB
.
Reproductive Physiology of Marsupials
.
Cambridge
:
Cambridge University
;
1987
.
29.
van den Broek
AJP
.
Entwicklung und Bau des Urogenital-Apparates der Beutler und dessen Verhältnis zu diesen Organen andrer Säuger und niederer Wirbeltiere
.
Morphologischen Jahrbuch
.
1910;
XLI
:
437
68
.
30.
Woolley
P
,
Webb
SJ
.
The penis of dasyurid marsupials
. In:
Stonehouse
B
,
Gilmore
D
, editors.
The Biology of Marsupials
.
London and Basingstoke
:
The Macmillan Press Ltd
;
1977
. p.
307
23
.
31.
Zug
GR
.
The penial morphology and the relationships of cryptodiran turtles
.
Occ Pap Mus Zool
.
1966
;
647
:
1
24
.
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