US ISSN 0006-9698

Cambridge, Mass. 16 January 2015 Number 540

REDESCRIPTION OF STICHOPATHES POURTALESI BROOK, 1889
(CNIDARIA: ANTHOZOA: ANTIPATHARIA: ANTIPATHIDAE)

Marzia Bo1 and Dennis M. Opresko2

Abstract. Stichopathes pourtalesi Brook, 1889, the type species of the genus Stichopathes Brook, is redescribed
on the basis of an examination of the type material originally identified by de Pourtales as Antipathes desbonni
Duchassaing and Michelotti and as Antipathes spiralis Pallas. Because the type series consists of multiple species, it
was necessary to select a lectotype to clearly define the species. The species is characterized by a spiral, unbranched
corallum; triangular, compressed spines, some of which have one or more lobes at the apex; very faint elongate
papillae on the surface of the spines extending from near the apex to as much as one-half the distance to the base; and
large polyps with long tentacles (in preserved specimens). The species is similar to Stichopathes gracilis (Gray);
however, scanning electron microscopy indicates that there are distinct differences between the two species in the
microscopic surface ornamentation of the spines.

Key words: black corals; morphology; taxonomy; syntypes; lectotype; paralectotypes

INTRODUCTION

In 1889 Brook established the genus
Stichopathes and included within it seven
species: S. pourtalesi Brook, 1889, S. gracilis
(Gray, 1857), S. echinulata Brook, 1889,
S. desbonni (Duchassaing and Michelotti,
1864), S. occidentals (Gray, 1860), S. filifor-
mis (Gray, 1868) and S. luetkeni Brook, 1889

1 Universita degli Studi di Genova, Genova, Italy; e-mail:
marzia.bo@unige.it.

2U.S. National Museum of Natural History, Smithso-
nian Institution, Washington, D.C., U.S.A.; e-mail:
dmopresko@hotmail.com.

(original spelling liitkeni). Brook did not
specifically state which of the seven species
he considered the type species of the genus;
however, in his description of the genus he
refers to information provided by de Pour-
tales on the polyps of specimens that de
Pourtales had identified in 1872 and 1874 as
Antipathes desbonni Duchassaing and Miche-
lotti and also those he identified in 1880 as
Antipathes spiralis Pallas, 1766. Brook (1889:
88) notes that he established the genus “in
consequence of what Pourtales says about
his Cirrhipathes spiralis (Pourt. non Pall.),
and in order to distinguish those unbranched

® The President and Fellows of Harvard College 2015.

?

B RE VI ORA

No. 540

forms ol Antipathinae having polyps on one
side ol the axis only, from those which, as in
the genus Cirripathes have them distributed
spirally all around the stem.” Brook (1889)
indicates that all of the other six species that
he referred to Stichopathes were represented
in the British Museum collection by speci-
mens not preserved in alcohol and for which
the polyps “were not available for study.”
For two of these species, S. echinulata and S.
gracilis, the polyps were described as being
distributed in a single row; however, Brook
provides no additional information. The
species that Brook discusses first in his
monograph is S. pourtalesi , and because the
type species was always the First species of a
genus discussed (e.g., in the case of Anti-
pathella , Aphanipcithes , Tylopathes, Schizo-
pathes, Bathypathes, Taxipathes, and Clado-
pathes), it must be concluded that he
considered S. pourtalesi the type species of
the genus. Because Brook’s establishment of
the genus Stichopathes was based solely on
de Pourtales’ description (Brook did not
examine any of the material de Pourtales
identified as A. desbonni or A. spiralis), and
because de Pourtales’ description is limited in
scope and was based on numerous specimens
representing multiple species, it is necessary
to redescribe and clearly define the species
and to select an appropriate lectotype.

MATERIAL AND METHODS

This report is based on specimens identi-
fied by de Pourtales in 1874 and 1878 as A.
(C.) desbonni (that he reassigned in 1880 to
A. spiralis), as well as additional specimens
collected during the “Blake” Expedition to
the Caribbean that de Pourtales identified in
his 1880 paper as A. spiralis. Almost all of de
Pourtales’ material is housed at the Museum
of Comparative Zoology (MCZ) at Harvard
University. At least three specimens are
known to be in the collections of the U.S.

National Museum of Natural History
(NMNH), Smithsonian Institution and one
sample in the collections of the Natural
History Museum, London (BM).

Photographs of the skeletal spines were
made using a scanning electron microscope
(SEM) housed at the NMNH. Analysis of
the morphometries of the skeletal spines was
conducted from direct examination of the
material using a low-power ocular micro-
scope or by direct examination of photo-
graphs taken with the SEM.

Abbreviations: MCZ (Museum of Com-
parative Zoology); NMNH (U.S. National
Museum of Natural History, Smithsonian
Institution); BM (The Natural History Mu-
seum, London); USNM (prefix for NMNH
catalogue numbers).

Stichopathes Brook, 1889

Type Species. Stichopathes pourtalesi
Brook, 1889

Diagnosis. Monopodial unbranched cor-
allum (rarely secondarily branched following
regeneration of the broken stem), stem
straight, wavy, curved, or partially (upper
part of stem) or entirely spiral. Polyps
confined to a regular, single row on one side
of axis. Spines triangular to conical, smooth
or papillose, and simple, bifurcated, or with
multiple knobs or lobes at apex.

Remarks. The family Antipathidae cur-
rently contains three genera of unbranched
antipatharians: Stichopathes Brook, 1889;
Cirrhipathes Blainville 1834; and Pseudocir-
rhipathes Bo & Bavestrello, 2009 (in Bo et al.,
2009). In Stichopathes and Pseudocirrhi-
pathes, the polyps tend to be confined mainly
to one side of the axis, whereas in Cirrhi-
pathes the polyps occur in multiple rows that
often cover about three-quarters of the
circumference of the axis, leaving only a
small portion free. Pseudocirrhipath.es is
distinguished from Stichopathes by the fact
that the polyps tend to be arranged in

2015

RE-DESCRIPTION OF STICHOPATHES POURTALESI

3

irregular rows on one side of the axis,
whereas those in Stichopathes are in a
straight, regular row (Bo et ah, 2012).

Stichopathes pourtalesi Brook, 1889
(Figures 1-5)

Syntypes. In his 1874 paper de Pourtales
indicates that material he identified as A.
deshonni was collected at one station off
Barbados, and in his 1878 paper he lists four
stations where material identified as the same
species had been collected off Sand Key,
Florida (see Table 1). In his 1880 paper, de
Pourtales states that the specimens he
previously identified as A. desbonni he now
considered as A. spiralis Pallas. He further
states that this same species was very
common, having been collected at 23 stations
in depths ranging from 45 to 878 fm off
Havana, Santa Cruz, Montserrat, Martini-
que, St. Vincent, the Grenadines, Grenada,
and Barbados. On the basis of an inventory
of antipatharians in the MCZ collection
dated June 1969, as well as of identifications
written on the original labels, and an
examination of material at the NMNH, the
specimens of the type series and the stations
at which they were collected were identified
(see Table 1).

It should be noted that 18 Blake stations
corresponding to de Pourtales’ 1880 publi-
cation are shown in Table 1, whereas de
Pourtales mentions that the species was
collected at 23 stations. Either the specimens
from five stations were lost or donated to
other institutions without records being kept,
or, more likely, de Pourtales was including,
in the total of 23 stations, the single station
mentioned in his 1874 paper and the four
listed in his 1878 paper, in which case all 23
stations would be accounted for. Of the
stations listed above, only the specimen from
Blake Sta. 249, listed in the 1969 MCZ
inventory, could not be found.

Although specimens in the type series were
collected at 23 stations, over the years,
samples from some of these stations were
subsorted, reidentified, and given separate
MCZ catalogue numbers (this information
was never published). Because some of the
lots consisted of broken pieces of colonies, it is
possible that parts of the same colony were
catalogued as separate specimens, and equally
possible that parts of different colonies were
catalogued under the same registry number.

In total, we analyzed 32 lots, and because
some contained more than one colony or
multiple fragments representing more than
one specimen, we identified 51 different
specimens, of which 20 have been assigned
to S. pourtalesi sensu stricto (Table 2). All of
the remaining specimens in the original type
series represent different species. Until a
complete taxonomic revision of the genus is
completed, no attempt will be made to assign
these specimens to specific taxa; however,
similarities to other nominal species will be
pointed out where possible.

de Pourtales Descriptions. In his 1874
description of A. desbonni , de Pourtales
reports that his specimens are 50-55 cm
long, not much more than 1 mm in diameter
at the base; densely covered with small
spines, and with polyps that are of the type
with long tentacles, confined to one side of
the stem and alternately large and small. In
his 1878 description he adds that the colonies
are always spiral, the spines are in six rows
(presumably counting those only visible in
one view), the polyps cover four of those
rows, and that the tentacles are large and
fleshy. In his 1880 paper he elaborates that
the spirals are 10 to 20 cm in diameter; can
be either right to left or the reverse, or
change from one to the other in the same
colony; that the spines are short, triangular,
compressed, and arranged in a quincunx;
and that the coensarc on the side of the stem
opposite the polyps shows transverse canals

Table 1 . Station data and museum catalogue numbers for the syntypes of Sticiiopa tiies pourtalesi.

B REV I ORA

No. 540

o

in

oo

VO

VO

VO

<N

VO

VO

>n

m

o

ON

N

U

2

m

ON

ci

o

o

in

ro

OO

in

m

o

ON

r~~

oo

ro

VO

vO

z

C<7

UN

CO

SO

o

o

T3

3"

i/~>

vri

NO

in

S

NO

CO

CN

CO

m

CO

co

v

3

CO

D

SO

CQ

o

T— •

o

o

o

(N

.O

o

SO

On

SO

OO

Os

ON

ON

SO

OO

4_

ON

—

*K

(N

r-

3"

o

<N

^ (N

CO

„

in

ON

oo

O co

ON

NO

SO

NO

3"

3"

3"

3"

»o

3"

m

m

3"

y—-

3"

3 vo

m

m

m

r<3

m

m

co

co

CO

S rn

co

CO

m

CO

co

co

co

co

o

o

O

o

o

O

O

o

o

o

o

o

o

o

E o

o

ON

os

ON

ON

ON

ON

ON

ON

Z os

ON

ON

os

ON

ON

z

ON

a on

ON

N

N

N

N

N

N

N

N

c^ i. -«

Bu

N

N

N

N

N

C/D

N

■| N

N

U

U

U

U

U

U

U

U

U

U

U

U

U

U

2 u

U

S

S

s

S

S

s

S

2

s

s

S

S

S

2

2

a 5

C/3

2

a

<u

Q

NO

o

/— N

/— « V

VO

* S

o

o

*— 1

i ^

v

m

V

(N

so

7f

1

1—1

OS

1—1

OS

so

oo

CN

m

^t

o

»— 1

OO

so

so

r-

o

r-

r-

OO

*— <

(N

»—

(N

m

1-^

(N

(N

i—i

i—i

■ ^

' — ^

's— y

s— ^

^ — '

^ — '

' — ^

Swc'

■ /

n>— ^

o

r-

o

r-

oo

oo

oo

o

oo

os

so

^t

7f

SO

o

oo

(N

o

oo

r-

oo

oo

(N

so

os

os

so

(N

OS

(N

■ ‘

oo

oo

’ — 1

(N

C/3

<D

c

o

o

U

c3

z

<n

Z

Z ©

m

(N

ON

OO

z °°

C-l o
lO OO
o OO

C IN

Z O

t <N
m 1+
T' £

m

r-

G-

so

G SO

G —

(N

SO

g

a

^

z

7f

z

Cn

z

O

z

co

z

uo

-

uo

z

SO

(N

T— <

z

i— ■

V-)

oo

O

IZS

Cn

o

r-

Cn

OS

O

sn

Co

o

»— —

<N

o

o

1—1

o

rn

o

oo

0

<N

o

CN

o

un

o

r-

o

<N

*— 1

CN

l— '

i—

1— «

»— «

(N

(N

o

o

SO

o

so

o

so

o

so

0

so

o

SO

0

so

SO

m

CN

1

m

03

O

o

_1

(U
J3

E «3

3 a3

Z Q

c -a
o c
\c 3
cd

CO

m

O

■a

cd

x>

u.

cd

ca

Lu

(U

u

73

C

3

CO

o

<U

-O

C/3

CO

cd

?| o w
• =P S « pj -a

co

cd

C

3

>

X

N

3

< C ^

3 cd

X

'5

• S u
00 w

G r-

a 3 „

co ■ - CO
I x*

<U

a

3

_o

<U

-a

3

3

o

J*S

C

3

CO

o

X)

3

CO

3

c

aj

C/3

c

O

G G

j-i

J-I J-H

d> d)

C/3 C/3

*-* +-*
3 3

O O

s s

d)

w

G

3

?

■ Z

’g

's

■4— »

* ) J

J-

J-I

cd

cd

d>

_G

'O

G

C

d>

u.

o

3 M 3
O o o
0 3 0
3 ■- 3

•— 73 ■ —

> S >

+j d> +j

c/d r: fe oo

G XT*

d) G
o -a

C G

C c

d)

C/3

d)

_C

•6

G

C

d>

•—

o

G

d>

o

C

£ o >

O

o

G

o

G

o

O

o

o

o

o

o

o

o

OS

o

OS

O

o

o

OS

C/D

OS

OS

Os

Os

r-

r-

r-

os

r-

r-

oo

oo

oo

r-

oo

oo

oo

oo

1

—

*— 1

oo

m

>.

>,

J-I

>Y

J-I

J-I

>>

so

J-I

J-I

j-i

j-i

cd

cd

G

i— ■

cd

cd

cd

cd

G

G

cd

cd

G

G

G

G

j-i

j— i

j-.

(N

G

G

cd

J

o

^T)

C

cd

*—>

so

G

cd

►—5

r-

oo

G

cd

►— i

m

j-i

X>

d)

U*

X>

d)

JO

d>

U-

o

ON

rn

X)

d>

Uh

ON

cd

3

so

cd

m

so

IT)

SO

»zs

so

Os

o

o

m

o

3"

r-

NO

G

3

m

G

m

*— 1

1

in

<N

(N

(N

(N

<N

(N

(N

(N

Os

OO

X

p

g

G (D

o o
\G G
- <L>

"2 ro

w ^

r-

^ — s

^ ^

^ S

^ V

c

oo

oo

OO

oo

oo

o

i— i

r-

r-

oo

oo

oo

oo

."G

c/T

1— H

—

1

73

d)

G

-4—*

C/3

C/3

C/3

C/3

a

<d>

'J)

X

W

J— 1

G

G

G

G

G

o

j-i

J-i

j-i

J-.

j-i

CL-

G

G

G

G

jj

O

O

O

O

<s>

C/3

<u

3

a*

Pu

CP

Pu

G

<l>

d)

d)

d>

73

3

7D

73

hC

o

oo

oo

3

O

■a

G

<u

j-.

a

G

X

o

W

CP

<D

d>

J*

3

_3

CQ

o

O

O

o

O

O

O'

o

O

O

O

O

oo

oo

oo

oo

OO

oo

oo

oo

OO

OO

oo

OO

oo

oo

oo

oo

oo

oo

oo

oo

oo

oo

oo

oo

•— 1

d-/

oo

I—*

1

1

1

4— '

1—1

r.

oo

,v

r.

,v

c^

C/3

c^

c/)

C/3

C/3

C/3

C/3

CZ)

c/3

c^

C/3

^aj

'QJ

'P

UD

^d>

uu

G

G

G

C/3

G

G

G

G

G

G

G

G

G

-4—*

-4—*

-4—*

■4—>

■4— »

4-*

J-I

J-I

J-I

J-I

Jh

j—

J-l

J-I

J-i

u

J-i

J-I

G

G

G

G

G

G

G

G

G

G

G

G

G

O

O

O

O

O

O

O

O

O

O

O

O

CP

CP

CP

G

CP

CP

CP

CP

CP

CP

CP

CP

CP

d)

d>

d>

O

d)

d>

d)

d)

<u

d>

d>

d)

d)

73

73

73

CP

73

73

73

73

73

73

73

73

73

Table 1. Continued.

5

2015

RE-DESCRIPTION OF STICHOPATHES POURTALESI

N o
oo

NO

VO

V3

o

oo

m

ro

O

On

oo

r-

On

NO

»/">

rn

CO

l/~!

c*~i

m

ro

CO

O

o

O

o

ON

ON

On

ON

N

N

N

N

U

U

u

U

s

2

S

S

E

I

X

Oh

1)

Q

CD

r-

m

r-

oo

4-

oo

o

(N

O

O

u

^

z ^

i o _
CN

-=g«a3~si

o

o

OS

i/D

m i. m

os

O

m

o

o

^t

r-

m

o

Os

UD

ctf

cj

o

hJ

C/3

o

C/3

O

C/3

o

T3

"O

c3

X

X

X

S-H

S-H

S-H

X

CO

CO

Ch—

<U->

4-h

C^

<4^

O

O

o

c/3

o

'O

03

X

S-H

c3

CQ

<D

X

S S

O r3

Z Q

C ~0
O C
\n cd
c3

5o

ON

ON

ON

ON

r-

r-

r-

r-

oo

oo

oo

oo

T— 1

i—

X

X

X

X

o

CJ

o

CJ

S-H

S-H

—

S-H

C3

<N

2

o

2

NO

2

OO

2

r-

ON

ON

O

ON

O

(N

i/D

(N

ON

<N

*— i

CN

o

CL

O

OO

oo

'ju

P

O

CO

O

O

OO

OO

oo

oo

' — 1

c n

c /f

'JJ

P

a

4— >

S-H

S-H

P

p

O

o

0-

Co

<U QJ <U

T3 T3 "O

<D

T3

between the polyps that are more transparent
than the rest of the tissue, de Pourtales
further notes that the largest specimen is
3.20 nr long and 4-5 mm in diameter at the
base. In his 1880 paper de Pourtales provides
illustrations of the polyps and spines of a
specimen he assigned to A. spiralis , although
the specimen was not specifically identified
(nor was the station at which it was
collected). On the basis of these illustrations
(see Figs. 1A, B), the larger polyps can be
estimated to be about 1.5 mm in transverse
diameter, with three occurring along 4.5 mm
of axis (a total of five or six polyps per
4.5 mm of axis, if smaller polyps between the
larger ones are included). The illustration of
the spines is of low magnification and low
resolution, and only a very rough estimate of
the size of the spines can be made. The
largest spines appear to be about 0.1 mm tall
and arranged in irregular rows, with five or
six rows visible. No surface features are
discernable, and de Pourtales does not
mention any in the text. There is no certainty
as to whether the illustrations of the polyps
and spines were based on samples taken from
the same specimen.

Designation of a Lectotype. The key
taxonomic features that Brook (1889) used
to establish the genus Stichopathes were the
arrangement of the polyps in a single row
and the very long digitifonn tentacles that de
Pourtales stated were characteristic of the
specimens he identified as A. spiralis. Brook
(1889) lists the de Pourtales 1880 paper first
in the synonymy of S. pourtalesi, and it is
only in the 1880 paper where de Pourtales
presents an illustration of the polyps. In
selecting a lectotype for S. pourtalesi the
primary consideration was to find a speci-
men from the type series whose polyps
conform to these illustrations, and also one
that was part of the material reported on by
de Pourtales in 1880. Of the specimens of the
type series examined, it was decided that one

Table 2. Main taxonomic characters for the syntypes identified as Sticiiopathes pourtalesi sensu stricto.

6

B REV I ORA

No. 540

u>

c

0)

G

53

CL

G

G

X

CJ

>

>

G

CJ

>

_X

X

c n

0

C/3

CJ

CJ

c«

S 6

Q 2

_c

0

53

O

CJ

C/3

O

»

O

0/3

0

-4—4

O

CJ

<u

O

■4—1

CD

C/3

Q-

u

six

>

c:

X

’3

£

X

C/3

>

&

six

>

G

>

G

>>

v

O

G

r—

CL

cn

G

r "•

E

E

E

c*

X

G

>

C/3

s — '

£

G

2

G

£

*Z)

0

vn

O

rp

0

vn

00

m

IZh

vo in

u

H

o3

5

O

O —

'O

s

C/3

'O

TD

T3

T3

T3

no

T3

G

LG

LG

IS

CJ

s

G

G

G

G

_G

Z

S

z

5

z

G

z

cl

C/3

0)

G

CJ

0

u

CJ

CJ

u

CJ

0

r-1

E

E

E

C/3

E

E

E

E

£

£

Q

G

0

0

0

CJ*'

0

CJ

0

0

CJ

0

0

0

o3

cn

C/3

C/3

t/3

03

on

C/3

0-1

zS

C/l

C/3

C/2

C /3

G

0:

CJ

C-T

cj"

4J

?0

cj"

<D

iS

CJ

aj

CJ

oj

CJ

G

<u

G G G

■4—*

CL

C/3

on

C/3

'5-

i/1

CL

on

C/3

G

'cl

on

c^

C/3

C/3

on

C/3

cn cn cn

O

z

G

CL

JD

JO

03

CL

_o

o3

p

JD

bX)

CJ

03

CL

O

_o

JD

O

_o

-2-2-2

£

‘CL

’5.

'5-

cZ

'5.

‘CL

a

u

u.

’a

'5-

’a

'5.

*5.

'5-

■5,

CL CL CL

03

03

03

o3

03

as

03

o3

03

o3

aj

o3 G

CL

CL

CL

D.

CL

CL

a

P.

CL

CL

CL

CL

a

CL CL CL

</3

£

zz

.2?

0)

G

CJ

o

CE

’53

X

>

G

>

CJ

X

on

_><

X X

0

3

on

to

on cn

o

O

O
■*— *

O

O

O

O O

g

G

Z

v-.

0

G

CJ

■\

CJ

0

CJ

S-H

0

<u

CJ

CJ

CJ

m aj

>

r>

CJ

cn

>

><

*

>

_><

X

>

>

CJ

C/D

>

X

_><

.X > >

<£

LG

£

cG

£

C/3

’on

G

G

LG

’on

C/3 G cG

(N

G

C/3

G

C

G

c

03

s—

03

?

VO

nt;

vq

vq

°C

00

00

vq

vq

an

m

m

rn

nf;

vq

a) vt ^

’cl

-4—*

C/3

CL

E

O

0

0

0

0

O

0

O

0

0

O

O

O

0

0

O

0

OOO

C/D

s

<

O

0

VO

00

0

vo

(N

00

0

0

O

00

vo

00

00

r~~

00 O O

p

,, — „

- — «

1

O

O

*— <

1

O

’ — j

O

i— 1

O

0

0

O

0

O — —

mi

E

O

0

O

0

0

0

O

O

0

0

O

O

O

0

0

O

0

OOO

'u

E

CN

0

CN

0

O

CN

vo

vo

O

fN

CN

0

(N

CN

0

vo VO

*— -

»— <

1

(N

(N

»— 1

»— ■

1

i—*

1

O

0

0

0

O

O

O

O

O

0

O

O

O

0

0

O

0

OOO

£

-4—*

Diam

'-"“v

GO

£

00

r-

0

O

iz^

r-

O

O

O

r-

VO

m

T^

oc

(N

C3

Ov rn

X

ca

§

a

O

0

O

oi

rn

(N

0

O

0

0

0

’ — '

cb

OOO

s

m

0

O

g

r-

0

m

O

Ol

VO

(N

VO

0

g

G

O

m

m

oc

m

r-

CN

m

m

at ON OO

o3

s — y

in

r— (

0"

Gv

vo

vo

’ — 1

(N

(N

’ — 1

' — 1

, .

-C

cj"

cj"

QJ

CJ

K

CJ

r.

CJ

CJ

- O <L>

r*

u.

*— >

X

X

X

X

X

cj"

Ut

X

c^'

u.

X

u>

u.

X

x .E .E

c

03

bX)

G

G

CJ

‘

CJ

CJ

'p

C/D

■0

CJ

■0

G

*0

CJ

■*—4

■0

0

G

CL

g

CL

CL

CL

c

CL

CL

G

o3

G

a

G

E

G

CL

G

G

cx

CX C c

g

G

03

03

CJ

03

03

CJ

-O

CJ

o3

CJ

G

o3

CJ

G

rt

cd u w

03

U*

H-l

o

u

C/3

C/3

C/3

C/3

C/3

C/J

C/3

C/3

C/J

0

C/D

C/3

Vi

■4—*

*

-4— »

CJ

0

/— »

c

c

c

G

c

G

G

G

C

c

c

CJ

CJ

CJ

CJ

CJ

CJ

CJ

CJ

G

CJ

u

<u

CL

£

E

E

£

E

E

E

E

bX)

G

L_,

E

E

E

03

ao

Ml

tj)

M)

M)

Ml

M)

MO

Ml

bX)

MO

-C

03

«S

C3

o3

ctf

o3

cG

o3

o3

03

C/D

Ut

i—

u.

,u.

,5—

,5-

,Uh

*

C-H

<4—

C4—

■4—*
<— ■

-4— »

0*— (

04— .

O4— < -4—* -4—*

fH

OX)

ZZ

OX)

TD

-O

TD

T3

"O

~o

T3

Z-.

bX)

Z—

bp

G

Z—

.2?

T3

G

T3 M M

JJ

JJ

JJ

^CJ

CJ

CJ

CJ

CJ

CJ

CJ

G

G

<U •— .—

2

2

"0

70

70

70

70

3

3

70

S3

70

70

" cS cS

j-.

i—

’o

‘5

’O

’0

O

O

O

O

O

i—

1—

O

j—

•4—4

O

O

0 E Z

C/3

C/D

CJ

CJ

CJ

CJ

CJ

CJ

CJ

CJ

CJ

C/3

on

CJ

c^

G

G

G cn cn

0

(^1

4—1

<N

m

«— 1

m

m

— CN

z

'W ^

S— ^

s— ^

s— ^

'N— y

s— ^

S— ^ 'N-^ s-x'

<D

m

r-

(^1

r^»

m

t—

00

rn

vn

vo

r-

r-

r-

r-

00

oc

00

00 — —

00

»Z3

vn

»Z3

«Zh

VT)

in vo vo

bX)

o

(N

m

m

m

m

m

m

m

m

m

m

m

ro

m

m

m

m m m

vo

vo

O

O

O

O

O

O

O

O

O

O

O

O

O

O

O

OOO

VO

vo

ov

ov

ov

OV

a\

OV

GV

ov

ov

GV

GV

GV

GV

GV

GV

GV GV GV

03

*—>

N

N

N

N

N

N

N

N

N

N

N

N

N

N

N

N

N

N N N

a

u

U

U

U

U

U

U

O

U

U

U

U

U

U

U

U

U

U

u u u

S

s

s

S

S

s

2

S

2

2

s

s

s

s

2 2

s

■Numbers in parentheses designate specific specimens in same lot as assigned by study authors.
2Height of spines is indicated for polypar/abpolypar sides of axis, respectively.

^Length of corallum is given for entire reconstructed colony or for indicated section only.

2015

RE-DESCRIPTION OF STICHOPATHES POURTALESI

7

from Blake Sta. 156 (MCZ 90342), although
not a complete specimen, had polyps almost
identical in size and appearance to those in
the 1880 publication (see Figs. 1C, D). This
specimen was selected as the lectotype of S.
pourtalesi Brook.

Diagnosis. Unbranched corallum, either
spiral from near base or with lower portion
relatively straight and upright and upper
portion forming large coils or spirals. Spirals
10 to 20 cm in diameter. Spines triangular to
conical, extending out at right angles to axis
or slightly inclined distally. Polypar spines
usually simple or with slight bifurcation at
apex; mostly 0.14-0.17 mm tall (maximum
about 0.2 mm); lateral spines in some places
with one or more irregular lobes at apex;
abpolypar spines simple, reduced in size.
Polypar and abpolypar spines with faint,
fusiform papillae. Papillae covering spine
surface from apex, or just below apex, one-
third to one-half distance to base. Spines on
thicker parts of stem more often multilobed
at tip, and papillae becoming more rounded
and distinct. Polyps up to about 1.7 mm in
transverse diameter, with long distally di-
rected tentacles; basal section of proximal
lateral tentacles extending distally around
mouth; proximal lateral tentacles up to
about 3 mm long; sagittal tentacles up to
5 mm long (in preserved material); four to
seven polyps per centimeter.

Description of Lectotype. The specimen
selected as the lectotype (MCZ 90342, Blake
Sta. 156) consists of three broken, curved
pieces of stem whose total length is 43 cm
(19 +14+10 cm) (Fig. 2). The diameter at
the lower end is about 0.7 mm. The spines on
the polyp side of the axis are compressed and
triangular in lateral view (Fig. 3). On a
section of stem 0.5 mm in diameter they are
up to 0.17 mm tall, and spaced 0.5-0. 7 mm
apart from each other, resulting in slightly
less than three per millimeter. They are
simple or with a slight bifurcation at the

apex and are orientated at right angles to the
axis. Spines occur in seven to eight longitu-
dinal rows (as seen in one view). The spines
in the rows lateral to the polypar spines show
more distinct bifurcations at the apex and
some are multilobed. The abpolypar spines,
those directly opposite the polypar spines,
are reduced in size, as small as 0.06 mm in
places, and simple — without bifurcations or
lobes at the apex. Fusiform papillae cover
part of the surface of the spines (Figs. 3D,
E). The largest papillae are up to about 10 pm
long and 2.5 pm wide, and in areas where
they are most abundant have a density of
about 40 per 1,000 pm2. They cover the
surface from just below the apex to one-third
to one-half the distance to the base. On some
spines the papillae become less distinct
toward the proximal edge of the spines.

The polyps are in a single row on one side of
the axis (Figs. 1C, D). They have a transverse
diameter up to about 1.5 mm. Small polyps
can be found between some of the larger ones.
Six polyps occur along 1 cm of axis. The
sagittal tentacles are up to about 3.8 mm long
and the proximal laterals about 3 mm long.
Traces of long sweeper tentacles are visible (up
to 5 mm long). The basal section of the
proximal lateral tentacles extend distally and
are wrapped around the oral cone.

Species Variation. The other 19 specimens
in the type series that we assign to S.
pourtalesi , sensu stricto, conform, in general,
to the diagnosis given above; however, a
certain degree of intraspecific variability is
recorded among these specimens (Table 2).
Colonies generally show loose coils, al-
though straight sections can be present,
especially in very young specimens or at the
base of the very long ones. The height of the
polypar spines ranges from 0.1 to 0.2 mm,
whereas that of abpolypar spines from 0.04
to 0.16 mm. The spines (absent in the basal-
most 1 to 2 cm of the stem) are in four to
eight longitudinal rows (one view) and within

8

B RE VI ORA

No. 540

Figure 1. S tichopathes pourtalesi Brook. A, B, reproduction of de Pourtales’ illustrations of the polyps of
“ Antipathes spiralis.” C, D, polyps of lectotype of S. pourtalesi (MCZ 90342).

Figure 2. Lectotype (MCZ 90342) of Stichopathes pourtalesi Brook.

2015

RE-DESCRIPTION OF STICHOPATHES POURTALESI

9

each row they are 0.3 to 0.8 mm apart.
Papillose spines are present in all specimens;
however, the density of the papillae is
variable. Bifid and lobate spines are not
common and are generally restricted to the
thicker portions of both juvenile (less than
20 cm high) and adult colonies. There is no
clear difference in height or shape of the spines
in colonies of different ages, although spines
are usually smaller and less commonly bifid in
juvenile specimens. The polyps are 0.6 to
1 .7 mm in transverse diameter (from proximal
side of proximal lateral tentacles to distal side
of distal lateral tentacles) and there are four to
seven polyps per centimeter. Sagittal tentacles
are up to 5 mm long (including ones that
appear to be sweeper tentacles).

The most complete of the larger specimens
assigned to S. pourtalesi is MCZ 90343 (from
Blake Sta. 157). This specimen is broken into
16 coiled pieces, the total length of which is
about 470 cm (although the larger, thicker
pieces clearly belong to the same colony, the
same could not conclusively be determined
for all of the smaller apical pieces). The basal
section is straight and 4.8 by 5.1 mm in
diameter at its lower end, but a holdfast is
not present. The uppermost broken piece is
91 cm long and about 0.9 mm in diameter at
its lower end, which is curved in the form of
a coil about 9 cm in diameter. Intermediate
sections form coils up to 20 cm in diameter.
On sections of stem ranging in thickness
from 0.34 to 1.1 mm the polypar spines are
0.10 to 0.14 mm tall, and the abpolypar
spines are about 0. 1 mm. Height of spines is
slightly greater toward the basal portion of
the stem. The spines are in four to eight
longitudinal rows (one view), and within
each row they are 0.5 to 0.6 mm apart. On
apical sections of the stem, the spines tend to
show fewer papillae than those of the
lectotype (Fig. 4). Spines on thicker sections
of the stem show more coarse papillae and
multiple lobes at the apex (Fig. 5). The

polyps are 1.2 to 1.4 mm in transverse
diameter and there are five to six polyps
per centimeter.

Taxonomic Comparisons. The species that
most closely resembles S. pourtalesi, as
defined above, is S. gracilis (Gray, 1857).
The original description of S. gracilis is
rather brief and consists solely of the
following: “slender, tapering, slightly spi-
nose.” The type was described in more detail
by Brook in 1889, who reported that the
colony was curved but not spiral, and that
the polyps were arranged in a single row. The
spines were described as triangular, com-
pressed, standing out at right angles to the
axis, and forked on the thinner parts of the
stem, the forked spines eventually splitting
into two separate spines on the thicker parts
of the axis where “the majority of the spines
are now simple, but some are short and
broad, having the apex divided into a
number of processes giving a serrate appear-
ance.” A re-examination of the type speci-
men in the Natural History Museum (BM
Reg. No. 23.1.16.1) revealed that the polyps
are about 1.5 to 2.0 mm in transverse
diameter with four to five per centimeter.
On a sample of the stem 0.64 mm in
diameter, the polypar spines are 0.22 mm
tall, and the abpolypar spines 0.12 mm tall
(Figs. 6A-C). Five to six longitudinal rows
are visible in one view, and the spines are
about 0.9 mm apart in each row, resulting in
only two spines per millimeter in each row.
The apex of the spines is round and smooth
or has small lobes. Numerous small papillae
cover the area from just below the apex to
the middle of the spines or slightly below the
middle (Figs. 6D, E). The papillae are
roundish to oval — in the latter case with
the longer of the axes extending in a
direction from the apex of the spine to the
base. The density of the papillae in the center
of the side of the spines was estimated to be
about 40 per 1 ,000 pm2.

10

BREVIORA

No. 540

Figure 3. Schizolectotype of Stichopathes pourtalesi Brook (from MCZ 90342). A-C, sections of stem showing
the occurrence of both bifid and simple spines. D, E, close-up view of two spines showing the surface papillae and
slight apical bifurcation.

2015

RE-DESCRIPTION OF STICHOPATHES POURTALESI

Figure 4. Schizoparalectotype of Stichopathes pourtalesi Brook (from MCZ 90343). A and C, two sections of the
stem showing the arrangement and size of the spines. B and D, single spines with faint papillae.

■uirl oos-

12

BREVIORA

No. 540

A

Figure 5. Schizoparalectotype of Stichopathes pourtalesi Brook (from MCZ 90343). A and C, arrangement of
spines along the thicker section of the stem. B and D. single spines with papillae and apical lobes.

2015

RE-DESCRIPTION OF STICHOPATHES POURTALESI

13

Figure 6. Schizoholotype of Stichopathes gracilis Gray (USNM 100363; from BM Reg. No. 23.1.16.10). A-C,
one section of stem shown at increasing magnification. D, E, individual spines showing surface papillae and small
apical knobs.

14

B REV 1 ORA

No. 540

Table 3. Reidentifications of the syntypes of Stichopatiies pourtalesi Brook.

Species/Group
S. pourtalesi sensu stricto

Stichopathes spp. group A

(Spines taller than in S. pourtalesi sensu stricto with
sparse to dense tubercles or minute papillae)

Stichopathes spp. group B

(Spines shorter than in S. pourtalesi sensu stricto with
distinct tubercles or minute papillae)
Aphanipathidae?

Syntypes1

MCZ 66253, MCZ 66387, MCZ 90342, MCZ 90343 (1),
MCZ 90343 (2), MCZ 90347, MCZ 90348, MCZ 90353,
MCZ 90355, MCZ 90356, MCZ 90357 (1), MCZ 90357
(2), MCZ 90357 (3), MCZ 90357 (4), MCZ 90358 (1),
MCZ 90358 (2), MCZ 90358 (3), MCZ 90358 (4), MCZ
90361 (1), MCZ 90361 (2)

MCZ 66251 (1), MCZ 66251(2), MCZ 66251(3); MCZ 66385

(1) , MCZ 66385 (2), MCZ 66385 (3), MCZ 66385 (4),
MCZ 66385 (5), MCZ 66386, MCZ 90340, MCZ 90341,
MCZ 90344, MCZ 90350, MCZ 90360 (1), MCZ 90360

(2) , MCZ 90360 (3), MCZ 90362, MCZ 90363, MCZ
90365 (1), MCZ 90365 (2), MCZ 90365 (3)

MCZ 90337, MCZ 90338, MCZ 90339, MCZ 90349, MCZ
90351, MCZ 90352, MCZ 90354 (1), MCZ 90354 (2),
MCZ 90359
MCZ 90364

‘Number in parentheses after MCZ catalogue number designates a specific specimen from that lot as assigned by
the study authors.

On the basis of the information given
above, S. pourtalesi differs from S. gracilis in
the maximum size of the polypar spines
(generally less than 0.17 mm vs. 0.22 mm),
the maximum size of the polyps (1.7 mm vs.
2.0 mm), the maximum density of the polyps
(seven per millimeter vs. five per millimeter),
and the shape of the papillae on the surface
of the spines (distinctly elongated in pourta-
lesi and round to slightly elongated in
gracilis). The shape of the papillae is
considered a key feature in separating the
two taxa, and illustrates the importance of
evaluating high-resolution images of individ-
ual spines.

Distribution. On the basis of the syntypes
assigned to S. pourtalesi, the species occurs in
the Caribbean Sea and Gulf of Mexico
(Montserrat, Yucatan Bank, Barbados, Mar-
tinique, Florida, St. Vincent, and Grena-
dines) at depths ranging from 82 to 299 m.

Remarks on Other Specimens in the
Type Series. Not all the specimens referred
by de Pourtales to A. spiralis Pallas conform
to the diagnosis and description given above.

At least three other species groups may be
represented in the type series (Table 3).
These are discussed below:

1. Syntypes with relatively tall spines with
papillae or distinct tubercles (see group A
in Table 3).

Colonies are typically coiled or spiral; the
coils are 6 or 7 cm in diameter. In the
most complete specimens, the lower
portion of the stem is straight, whereas
the upper part is twisted in loose coils.
The polyps up to 2.5 mm in transverse
diameter, interpolypar space 0.8 to
1.2 mm, with three to four polyps per
centimeter and sagittal tentacles up to
2.4 mm long. Polypar spines up to
0.32 mm tall (as measured from apex of
spine through the center to the base);
abpolypar spines one-half to two-thirds
size of polypar spines; mutual distance of
spines mostly 0.9 to 1 mm; number of
rows usually five or six. In almost all
specimens, distinct apical tubercles are
present on spines from all regions of the

2015

RE-DESCRIPTION OF STICHOPATHES POURTALESI

15

stem: apex, mid-portion, and base
(Figs. 7, 8C).

A comparison with other nominal species
of Stichopathes indicates that these spec-
imens are related to two other nominal
species of Stichopathes: S. paucispina
Brook, on the basis of the expanded
description of the latter species by
Opresko and Genin (1990), and S.
dissimilis Roule (1902, see also Roule,
1905). In S. paucispina the polypar spines
are up to 0.32 mm tall and the abpolypar
spines up to about 0.24 mm. Both the
polypar and abpolypar spines have dis-
tinct tubercles (12 to about 17 seen in one
lateral view of a single spine) near the
apex. Polyps occur in two sizes, arranged
in a single, straight row. The larger
polyps are 2.3 to 3.4 mm in transverse
diameter and the smaller polyps 0.8 to
1.2 mm, with 2.5 to three polyps occur-
ring along 1 cm of axis. Sagittal tentacles
up to 4 mm long in larger polyps, usually
slightly longer than proximal lateral
tentacles; distal lateral tentacles up to
2 mm in the largest polyps.

Also included in group A (Table 3) are
specimens with tall spines that have only
a small number of tubercles (Fig. 8B), or
with surface features that resemble small
papillae rather than distinct tubercles
(Fig. 8A). Further study is needed to
determine if these forms represent differ-
ent species or only variants of the species
mentioned above.

2. Syntypes with small polyps and extreme-
ly small spines (see Table 3 and Figs. 8D,
E). All specimens form thin, coiled
colonies. Spines are simple, triangular,
and subequal (0.04-0.1 mm tall on the
polypar side and 0.02-0.06 mm on the
abpolypar side); arranged in four to five
longitudinal rows (one view); and are
0.2-0. 6 mm apart in each row. Spines are
slightly (Fig. 8D) to distinctly papillose

(Fig. 8E) over one-third to two-thirds of
the apical surface and, in numerous
specimens, are arranged in distinct verti-
cils along the apical or central portion ol
the stem. Polyps are around 1 mm in
transverse diameter, and are variably
spaced with five to nine polyps per
centimeter (the poor state of preservation
of the soft tissues in many specimens
does not allow precise measurements).
Polypar density tends to decrease toward
the apex of the stem.

3. Syntype with extremely tall spines
(Fig. 8F): MCZ 90364. One specimen in
three pieces totaling about 13 cm, with no
soft tissue remaining. Spines 0.52-
0.65 mm; somewhat club-shaped, with
very papillose surface; subequal in length.
A basal holdfast was not clearly present,
but a slight expansion of the sclerenchyme
at the thickest end suggests that the
specimen might have been a branch or
pinnule from a branched or pinnulated
colony. The size and shape of the spines
suggest that the specimen might belong to
a species in the family Aphanipathidae.

ACKNOWLEDGMENTS

We thank A. Baldinger of the MCZ for the
loan of the type material of Stichopathes
pourtalesi , and Shelia Halsey of the BM for
the donation of material of S. gracilis to the
NMNH. The authors also thank S. Cairns,
W. Keel, C. Bright, W. Moser, and P.
Greenhall for their assistance during visits
to the NMNH, and the two anonymous
reviewers of the manuscript. The photomi-
crographs were prepared in the SEM Labo-
ratory of the NMNH, Smithsonian Institu-
tion. S. D. Whittaker of the NMNH kindly
assisted in the SEM analysis. D. M. O. is a
Research Associate of the NMNH, and
gratefully acknowledges that affiliation.
Funding for this project was provided, in

16

BREVIORA

No. 540

Figure 7. Stichopathes sp. (syntype of S. pourtalesi Brook) from Blake Sta. 131 (MCZ 90344). A-C, sections of
stem of increasing diameter. D, E, single spines showing close-up view of surface tubercles.

2015

RE-DESCRIPTION OF STICHOPATHES POURTALESI

17

Figure 8. Stichopathes spp. (syntypes of S. pourtalesi Brook). A, single polypar spine from MCZ 90350. B. single
polypar spine from MCZ 90362. C, single polypar spine from MCZ 66385. D, single polypar spine from MCZ 90337.
E, single polypar spine from MCZ 90354. F, several polypar spines from MCZ 90364.

18

BREVIORA

No. 540

part, by a U.S. Department of Justice (DoJ)
grant to the Smithsonian Institution. The
authors are appreciative of the DoJ’s support
of black coral research at the Smithsonian.

LITERATURE CITED

Bo, M., M. Barucca, M. A. Biscorri, A. Canapa, H. F. N.
Lapian, E. Olmo, and G. Bavestrello. 2009.
Description of Pseudocinhipathes (Cnidaria: An-
thozoa: Hexacorallia: Antipathidae), a new genus
of whip black corals from the Indo-Pacific.
Italian Journal of Zoology 76: 392M02.

Bo M., G. Bavestrello, M. Barucca, D. M. Makape-
dua, A. Poliseno, A. Canapa, E. Olmo. 2012.
Morphological and molecular characterization of
the problematic whip black coral genus Sticho-
pathes (Hexacorallia: Antipatharia) from Indonesia
(North Sulawesi, Celebes Sea). Zoological Journal
of the Linnaean Society 166: 1-13.

Brook, G. 1889. Report on the Antipatharia collected
by H. M. S. Challenger during the years 1873-1876.
Report on the Scientific Results of the Voyage of
H.M.S. Challenger, Zoology 32: 1-222.
de Blainville, H. M. D. 1834. Manuel d’Actinologie on
de Zoophytologie. Paris, F. G. Levrault.
de Pourtales, L. F. 1874. Crinoids and corals.
Zoological results of the Hassler Expedition, III.
Catalogue of the Museum of Comparative Zoology ,
Harvard 8: 25-50.

de Pourtales, L. F. 1878. Report on the corals of the
Blake expedition. Bulletin of the Museum of
Comparative Zoology, Harvard 5: 197-212.
de Pourtales, L. F. 1880. Zoological results of the
Blake expedition to the Caribbean Sea. Bulletin of

the Museum of Comparative Zoology, Harvard 6:
113-118.

Duchassaing, P., and J. Michelotti. 1864. Supplement
au memoire sur les coralliaires des Antilles.
Memorie della Reale Accademia delle Scienze di
Torino (2) 23: 97-206, 10 pis.

Ehrenberg, C. G. 1834. Die Corallenthiere des Rothen
Meeres. Berlin, Gedruckt in der Druckerei der K.
Akademie der Wissenschaften.

Gray, J. E. 1857. Synopsis of the families and genera of
axiferous zoophytes or barked corals. Proceedings
of the Zoological Society of London 25: 278-294.

Gray, J. E. 1860. Notice of new corals from Madeira
discovered by J.Y. Johnson, Esq. Annals of the
Magazine of Natural History (Series 3) 6: 31 1.

Gray, J. E. 1868. Descriptions of some new genera and
species of alcyonid corals in the British Museum.
Annals of the Magazine of Natural History (Series 4)
2; 441 M45.

Opresko, D. M., and A. Genin. 1990. A new species of
antipatharian (Cnidaria: Anthozoa) from the sea-
mounts in the eastern North Pacific. Bulletin of
Marine Science 46: 301-310.

Pallas, P. S. 1766. Elenchus Zoophytorum Sistens
Generum Adumbrationes Generaliores et Specierum
Cognitarum Succinctas Descriptions cum Selectis
Auctorum Synonymis. Hagae-Comitum.

Roule, L. 1902. Notice preliminaire sur les Antipathaire
provenant des collections du Prince du Monaco.
Memoir es de la Societe zoologique de France 15:
328-339.

Roule, L. 1905. Description des Antipathaires et Cer-
ianthaires recueillis par S.A.S. le Prince de Monaco
dans l’Atlantique Nord. Resultats des Campagnes
Scientifiques du Prince de Monaco , Fasc XXX:
99 pp. •