MAUREEN E. DOWN Zjorocallida, New Order,

and Doraster constellatus,

New Genus and Species,

with Notes on the

^oroaster idae

Echinodermata: Asteroidea) A

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • 1970 NUMBER 64
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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 64

Maureen E. Downey Zorocallida, Ne w Order,

and Doraster constellatus,

New Genus and Species,

with Notes on the

Zoroasteridae

(Echinodermata:

Asteroidea)

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

1970
ABSTRACT

Downey, Maureen E. Zorocallida, New Order, and Doraster constellatus, New

Genus and Species, with Notes on the Zoroasteridae (Echinodermata: Asteroidea).

Smithsonian Contributions to Zoology, 64: 1-18. 1970.—During a study of the

Asteroidea of the Caribbean and Gulf of Mexico, a new genus of the family Zoroasteridae was discovered. A general survey of the family revealed apparent relationships

with the fossil Calliasterellidae and prompted the conclusion that both of these families

were incorrectly placed in the order Forcipulatida. A new order, the Zorocallida, is

therefore erected for the Zoroasteridae and Calliasterellidae. The new genus, Doraster,

is described, with the type-species D. constellatus. Several other observations on the

classification of the Zoroasteridae are made, including the synonymizing of three species

of Zoroaster found in the Atlantic.

Official publication date is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year.

UNITED STATES GOVERNMENT PRINTING OFFICE

WASHINGTON : 1970

For sale by the Superintendent of Documents, U.S. Government Printing Office

Washington, D.C. 20402 - Price 30 cents (paper cover)
Maureen E. Downey ZorOCallida, New Order,

and Doraster constellatus,

New Genus and Species,

with Notes on the

Zoroasteridae

(Echinodermata: Asteroidea)

As every asteroid systematist realizes, the order Forcipulatida has been a catchall for starfishes which could

not conveniently be placed in other orders. An artificial group, of obviously polyphyletic origin, the forcipulates should be accorded more attention than they

have received from systematists concerned with the

present unsatisfactory classification of the Asteroidea.

Tortonese's (1958) new order, Euclasteroidea, erected

to accommodate those manifest misfits, the Brisingidae,

was a decided advance toward the reorganization of

the Forcipulatida. It is hoped that the removal of the

Zoroasteridae and Calliasterellidae from this order,

proposed in this paper, will further elucidate the

definition of the f orcipulates. The Zoroasteridae are the

only living family heretofore referred to this order

which do not have crossed pedicellariae; whose skeleton does not form a reticulate network; and in which

the mouth plates are deeply sunken in the actinostome.

More important, they are also the only ones with

superambulacral plates—a very unlikely character for

a supposedly advanced group like the Forcipulatida.

Their removal leaves only the Heliasteridae and the

very large polyphyletic family Asteriidae (and probably some of the fossil Uractinina) in the forcipulates.

With Euclasterida and Zorocallida removed, the

way should be clear for a resorting and reassessment

Maureen E. Downey, Department of Invertebrate Zoology,

National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

of the Asteriidae and a better understanding of their

relationships.

The Forcipulatida have been defined as follows:

Pedicellariae of basal piece and two valves, either

straight or crossed.

Skeleton reticulate, forming a network of rectangular or very irregular meshes.

Podia in two or four rows, always with a suckered

disc; with simple ampullae.

Papulae on all surfaces.

Paxillae never present.

Spines and tubercles not occurring in groups or

bundles.

Madreporite always on aboral surface.

Median carina frequently present.

No conspicuous marginals.

Aboral ossicles usually arranged in regular rows.

Adambulacral plates generally short and crowded.

Mouth plates frequently inconspicuous and sunken

in actinostome; with ambulacral plates prominent except in Brisingidae.

Generally stellate, with small disc and long tapering

arms; arms five to many.

Most of the statements in the above definition, drawn

from a number of sources (Fisher, 1911, 1928; Hyman,

1955; Spencer and Wright, 1966), are either qualified

in some way, or totally untrue for some asteroids heretofore included in the order. Table 1 shows the more

simple and straightforward definitions which result

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 1.—Comparison of major characteristics of Euclasterida, Zorocallida, and Forcipulatida

Characteristic

Pedicellariae

Skeleton

Podia

Papulae

Paxillae

Spines and tubercles

Median carina

Marginals

Aboral ossicles

Adambulacral plates

Mouth plates

Arms

Euclasterida

Always crossed.

Weakly developed, not

reticulate, of thin plates or

transverse arches widely

separated.

In two rows, with large suckered

disc; ampullae simple.

Wanting or confined to aboral

disc and arm bases.

Never present.

Singly, never in groups of

bundles.

Never present.

Never present.

In irregular transverse arches.

Not compressed.

Expanded, neither ambs nor

adambs predominant; not

sunken in actinostome.

Always more than five.

Zorocallida

Always straight.

Very regular longitudinal and

transverse rows of imbricating

plates.

In four rows, becoming two

distally; with large suckered

disc proximally, pointed

distally; ampullae bilobed.

None below inferomarginals.

Present only in certain fossil

forms.

Same.

Always present.

Present and well-defined.

In regular longitudinal and

transverse rows.

Compressed.

Adambs project into sunken

actinostome.

Never more than five.

Forcipulatida

Straight or crossed.

Reticulate, more or less open

meshwork.

In two or four rows, with large

or small suckered disc;

ampullae simple.

On all surfaces.

Never present.

Same.

Sometimes present.

If present, inconspicuous,

mostly irregular.

Mostly irregular.

Compressed.

Ambs predominant; sometimes,

sunken in actinostome.

Five to many.

from separating the Euclasterida and Zorocallida

from the remaining Forcipulatida.

ZOROCALLIDA, new order

DIAGNOSIS.—Disc more or less domed, with a definite

arrangement of enlarged primary plates; median carina on arms, with all arm plates in rather compact

and imbricating longitudinal and transverse series;

mouth-frame deeply sunken in actinostome; alternating

carinate and noncarinate adambulacral plates; at

least some tubefeet large and conical, with tiny suckered

disc.

CONTENTS.—Families Zoroasteridae (recent) and

Calliasterellidae (fossil).

DISCUSSION.—A new order, the Zorocallida, is here

proposed to accommodate the Paleozoic-Mesozoic family Calliasterellidae and the recent family Zoroasteridae. This new order suggests a common root with the

Forcipulatida and certain of the Valvatida. Like the

forcipulates, the Zorocallida have pedicellariae consisting of a basal piece and two valves. They have both

the quadriserial and biserial tubefoot arrangement,

while the forcipulates may have either arrangement.

In both orders, the adambulacral plates are compressed, and spines and tubercles occur singly, never

in groups or bundles. A median carina, always present

in the Zorocallida, is present also in some Forcipulatida. The Zorocallida share with many of the Valvatida

the orderly arrangement of primary disc plates, compact longitudinal and transverse series of arm plates,

similar mouth-frame, and the presence of superambulacral plates. It seems possible to theorize that the

Zorocallida are more ancient than the Valvatida and

Forcipulatida, with which they share certain basic

characters, and perhaps they may provide a clue to

the descent of the valvate and forcipulate starfishes

from a common origin.

The strong resemblance between the Zoroasteridae

and the Middle Devonian-Upper Cretaceous family

Calliasterellidae was noted by C. Wyville Thomson

(1873), the author of the first species described in

the Zoroasteridae. In his decription of Zoroaster fulgens, he called attention to the affinities of this species
NUMBER 64

FIGURE 1.—a, Calliasterella mira (Trautschold). Note the so-called "odontophore" displaced

toward the right, and the space to the left of it b, Calliasterella mira. Note the paired carinate

and noncarinate adambulacral plates. (From Schondorf, 1910.)

with Arthraster dixoni Forbes, an Upper Cretaceous

starfish from the lower chalk of Balcome pit near

Amberley, Sussex. Examination of Doraster, the new

genus described herein, confirms beyond a doubt that

these two families are indeed very closely related;

Doraster corresponds almost plate for plate with Calliasterella mira (Trautschold) in dorsal aspect (the

only one exposed with plates in situ), as far as can be

determined from photographs published by Schondorf

in 1909 (Figures la, b, 2a), and from a plaster cast

of part of the type, obtained through the courtesy of

Dr. A. N. Soloviev of the Paleontological Institute in

Moscow. I hestitate to place these two asteroids in the

same subordinate taxon, however, especially as Schondorfs careful and detailed study does not agree in

certain details from my own observations of his photographs and of Doraster. For example, Schondorf s reconstruction of the mouth-frame of C. mira is mostly

imaginary, as the mouth is not fully exposed in the

material he worked with. In fact, Schondorf himself,

in his discussion of the mouth-frame of C. mira, implies that his reconstruction is based on Viguier*s

(1878) figure of the mouth-frame of Asterias glacialis

( =Marthasterias glacialis), an assumption he justifies

by certain similarities between the ambulacral plates

of C. mira and the ambulacral pieces of the mouthframe figured by Viguier.

The alternating carinate and noncarinate adambulacral plates and the large conical tubefeet with small

suckered discs are characteristics unknown in the fossil

Calliasterellidae, but their presence can be inferred

from a reexamination of Schondorfs photographs of

C. mira. Scattered in the matrix of the specimen are

numerous, mostly dissociated, plates. Among them

Schondorf quite correctly identified several large carinate plates as adambulacrals, but he failed to notice

that, in nearly every case, a smaller noncarinate plate

lay next to them (Figure 16). In view of the condition

of this character in the Zoroasteridae, I believe these

smaller noncarinate plates are also adambulacrals

which alternate with the larger carinate adambulacrals,

just as in the recent Zoroasteridae. Given these simi-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 2.—a, Schondorf s reconstruction of Calliasterella mira. b. Schondorf s reconstruction

of cross section of arm of Calliasterella mira; compare with c, cross section of arm of Doraster

consttllatus.

larities, it is only reasonable to suppose that the tubefeet were also similar (although it is unlikely that we

shall ever know with certainty), especially in view

of the postulated relationship with the Valvatida.

As my experience with fossil asteroids is limited, it

would be best not to get deeply involved in a discussion of the similarities between other Calliasterellidae

and Zoroasteridae, but a few further comments on the

striking resemblance between Calliasterella mira and

Doraster seem called for. Schondorfs reconstruction

of C. mira shows a single, centrally located interradial

plate at the edge of the disc (Figure 2a), which he

calls an odontophore; however, careful examination

of his photographs shows this plate is actually displaced to the right of the interradial center (Figure

la), and there is a hiatus between it and the next plate

to the left, thus indicating the possibility that there were

originally a pair of plates in this position, as is the case

with Doraster. I also question Schondorfs interpretation of what he calls "spines" below the adradial

plates on the sides of the arms; his description of them

sounds more like small actinolateral plates, and again,

the photographs (Figure la) help to support this view.

Thus the reconstruction of the cross section of the arm

(Figure 26) by Schondorf is probably incorrect.

Incidentally, the species described by Kesling and

Strimple (1966) as Calliasterella americana, while

superficially resembling C. mira, seems a much more

"primitive" animal; the so-called adambulacral plates

are completely different, and there are definitely no

small plates between the "adradials" and the "adambulacrals." Also, the type and placement of the spines

is completely different.

Characters of the Recent Zoroasteridae

Enlarged disc plates, in a definite arrangement of

primary radials, interradials, central, and perhaps

additional plates, is a very important character in this

family, as in the order. So are carinal plates along the
NUMBER 64

mid-arm and longitudinal and transverse series of arm

plates, in most cases decreasing in number of longi-rows

from proximal to distal.

Alternate carinate and noncarinate adambulacral

plates are present in all recent Zoroasteridae, if we

eliminate the very doubtful genus Prognaster. (See

page 14.)

Superambulacral plates are present in all members

of this family, although they are present only proximally in most genera, and in Zoroaster are much

reduced.

Large, duck-billed pedicellariae are a distinctive

feature of many of the recent Zoroasteridae. No other

family has these huge curved pedicellariae, with one

valve longer and broad-topped. They have a decidedly

bird-headed look, more like avicularia than pedicellariae. They are very large and numerous in Dor aster,

slightly smaller and less curved in Zoroaster, shorter

and heavier in Myxoderma, like Doraster in Cnemidaster (the genus perhaps nearest to Doraster in many

ways), blunter and more equal-valved in Bythiolophus,

and frequently absent in Mammaster. Small straight

pedicellariae are numerous in all these genera. There

are no pedicellariae in Pholidaster, and they are unknown in Prognaster (if, indeed, Prognaster exists).

Tubefeet, changing from straight with a large

suckered disc within the peristomial cavity to stout

and conical with a very small suckered disc, in

four rows proximally, becoming two rows distally, are

another prominent characteristic shared by all living

Zoroasteridae.

All the members of this family are regularly fiverayed, with long slender, tapering arms, and a relatively

small disc.

The bathymetric range of the family is from 100 to

2000 fathoms, and they extend from about 50° N to

about 40° S, with the greatest number of species

occurring in the tropical Indo-Pacific.

Key to the family ZOROASTERIDAE

(Modified from H. L. Clark, 1920)

A. Disc plates conspicuously enlarged, more or less convex, bare, or at least not covered with

spinules; disc and upper part of arms skin-covered.

B. Disc plates distinctly stellate, not much swollen Dortuttr

BB. Disc plates decidedly swollen, more round than stellate.

C. Adradials absent; 3 rows of actinolateral plates Mammasttr

CC. Adradials present; 4-5 rows of actinolateral plates CnemidasUtr

AA. Disc plates plane, or, if convex, closely covered with spinules; no covering of skin on dorsal

surface.

B. Carinals smooth and bare; surrounded by series of skin-covered squamules; no

pedicellariae Pholidaster

BB. Carinals not surrounded by series of skin-covered squamules; both small straight

and large duck-billed pedicellariae present.

C. First superambulacral plate modified to form a conspicuous buttress connecting first two ambulacrals with body wall.

D. Adradials present; superomarginals not over lapping carinals

Myxodtrma

DD. Adradials absent; superomarginals strongly overlapping carinals

Bytkiolophus

CC. First superambulacral plate not modified to form a conspicuous buttress between ambulacrals and body wall Zoroastsr

Doraster, new genus

With the characters of the type-species, D. constellatus,

new species.

ETYMOLOGY.—Greek, dora=skin, hide, and aster=

star, (m.); specific name: Latin, con-stellatus = with

stars, starry (m.).

Doraster constellatus, new species

FIGURES 3-11

DESCRIPTION.—The plates of the disc dorsum are

large, flat, stellate, smooth, skin-covered. The central plate usually bears a stubby tubercle and a similar
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

t

FIGURE 3.—Dor aster constellates, side view (interradius). Note fused interradial adradials.

Holotype (USNM El 1352).

FIGURE 4.—Doraster constellatus (specimen cleared with chlorox). Small supplementary

platelets visible between carinal and adradial plates. Para type (USNM El 1353).
NUMBER 64

TABLE 2.—Measurements of specimens of Doraster constellatus

Specimen

Oregon 3583

Alaminos 21/

68-A-13

Oregon 382 A

B

C

D

£

F

Oregon II 10602

Albatross 2376 A

B

C

D

Oregon 2780 A

B

Albatross 2376 A

B

C

D

Albatross 2377 A

B

Albatross 2396

Albatross 2376 A

B

Albatross?

Preservation

dry

dry

dry

dry

dry

dry

dry

dry

dry

dry

dry

dry

dry

dry

dry

ale

ale

ale

ale

ale

ale

ale

ale

ale

ale

Major radius

R(mm)

80

121

122

*

105

*

95

110

183

66

119

109

118

239

*

180

134

110

87

134

136

57

66

46

102

Minor radius

r(mm)

12

15

12

14

11

11

13

12

23

9

11

16

16

38

18

17

21

13

12

18

17

9

9

8

15

Height of

disc (mm)

9

10

12

14

11

11

12

11

23

11

20

16

20

31

13

22

22

14

14

17

21

7

11.5

7

18

Height of

arm at third

carina (mm)

6

9

7

11

9

8

8

8

14

7

10

10

15

20

11

14

11

9

7

11

16

5

6

5

10

Number of

interradial

actinolateral

rows

5

4-5

3

3

3

3

3

3

5

2

3

3

3

4

4

3

4

3

+

+

+

2

+

+

3

Number of

terminal

actinolateral

rows

2*

1

«

*

0

*

0

0

0

0

0

0

1

0

0

0

0

0

+

+

+

0

+

+

0

Number of

carinals

52*

73

69

*

80

*

60

81

99

50

73

60

72

100

*

+

+

+

+

+

+

+

+

+

+

*A11 arms broken or regenerating.

+

Skin-covering obscures plates.

tubercle sometimes occurs on other primary disc plates.

The anus, surrounded by small spinelets, is located in

one of the angles of the central plate. Surrounding the

centrodorsum are 5 primary interradial plates, and

between and underneath them are 5 radial plates on

which the 5 pointed lobes of the centrodorsum rest;

just outside and partly overlaying the primary

interradials are 5 large primary radials; a pair of

interradial plates, entirely concealed by skin and the

overlapping primary radials, lie between and beneath

the primary radials and 4 enlarged imbricating (in

large specimens, fused) adradials (Figure 3) which

occupy the interradial angle. Small pedicellariae and

tiny granuliform platelets (Figure 4) occur between

the plates of the disc. The madreporite is slightly

smaller than the disc plates, raised, covered with

irregular channels, and located between a primary

378-414 O—70 2

interradial and the enlarged interradial adradials; it

actually rests on a pair of concealed interradials. The

first carinal is enlarged, about the same size as the

interradial adradials.

The interradial gonads are in grape-like clusters,

short, hardly extending into the arm; they are attached

interadially 1 on either side of the enlarged 1st superambulacral plates.

The jaw (Figures 5, 6) is one solid piece (made up of

2 pairs of adambulacral plates), projecting into the

peristome. Behind it are 4 pairs of adambulacrals joined

across the interradius by a tooth-and-socket arrangement strongly reminiscent of the way the ambulacral

plates are joined at the top of the ambulacral groove.

Internally, the front of the jaw is flaired like the carinate adambulacral plates and bears, on each side, 3

subequal spines completely covered with tiny straight
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 5.—Doraster constellatus, oral view from outside. Paratype (USNM El 1353).

pedicellariae, which thus form a wreath around the

peristome. External to these 3 pairs of oral spines is a

single pair of long sharp spines. The next 2 pairs of

adambulacral plates behind the jaw bear each a single

pair of long sharp spines, and the 2 pairs behind them

bear 4 to 6 similar spines. The 7th and 8th pairs of

interradial adambulacrals are separated by a large

duck-billed pedicellaria. The first 2 pairs of ambulacral

plates are also strongly fused and greatly swollen. The

ambulacrals curve sharply upward under the disc and

thus support the disc in a domed position. The 2 fused

pairs are buttressed against the body wall by a solid

ridge formed of the first 2 (enlarged) superambulacral

plates. A pore for the oral tubefoot occurs in the angle

between the fused ambulacrals and the buttressing

superambulacrals. Superambulacral plates of two

kinds occur, alternately, but do not continue to the

end of the arm; rather, they diminish in size and

eventually disappear. A straight, triangular superambulacral plate alternates with a thin, flaired superambulacral plate which terminates in a broad flat disc

resting on the ambulacral plates. They gradually become similar, small, and granuliform, and do not continue much beyond the middle of the arm. The tubefeet are in 4 rows proximally, 2 rows distally. Around

the mouth, they are straight and conspicuously annulated and terminate in a large suckered disc, but

beyond the confines of the disc, they are stout and

conical and taper to a tiny disc, indicating that the

original, more "primitive" condition in this order was

with a large suckered disc, while the pointed tubefoot

is a functional adaption.

On the arms, the carinal plates and 2 rows of plates

on either side (adradials and superomarginals) are

covered with skin, devoid of spines or spinules, but may

have small pedicellariae, especially proximally. The

carinals may have a central, non-spine-bearing tubercle.

Adradial plates overlap carinals and superomarginals.
NUMBER 6 4

FIGURE 6.—Doraster constellatus, oral view from within disc. This specimen shows also the skin

covering the dorsal surface of the arms, and the wreath of small pediccllariae around the mouth.

Paratype (USNM 10742).

Carinals and adradials about equal in size, somewhat

cruciform, wider than long; superomarginals slightly

smaller, lobed or, in large specimens, somewhat triangular, tapering toward actinal surface. On large specimens, an irregular row of tiny granuliform platelets

(Figure 4) occurs between the carinals and the adradials; a few also occur sporadically on the disc.

Proximally, there are 5 rows of plates between the

superomarginals and the adambulacrals covered with

small sacculate spinules; the row adjoining the superomarginals is the inferomarginal row. About halfway

out on the arm, one row of plates between the inferomarginals and the adambulacrals drops out, then further out, another, until at the end of the arm only the

carinals, adradials (by this time very small), superomarginals, and inferomarginals remain. The row of

plates adjoining the adambulacrals is very narrow,

longer than broad, and only the edge of the plate may

be visible. Most of the actinolaterals carry a long, flattened, appressed spine, directed upward and attached

to a prominent tubercle. All of these arm plates are in

regular longitudinal and transverse rows; however, the

latter do not correspond to the adambulacrals. Papulae

occur singly or in groups of up to 5 between the plates;

none occur below the inferomarginals. The terminal

ocular plates are cordiform, covering the arm tip, with

3 or more coarse terminal spines.

Adambulacral plates are of two alternating sorts,

carinate and noncarinate (Figure 7) ; the carinate

adambulacrals project strongly into the ambulacral

groove and bear 4 movable spines on a transverse row

of tubercles. The spines are stout and rounded at the
FIGURE 7.—Doraster constellatns, cleaned specimen, showing the carinate and noncarinate

adambulacral plates. Paratype (USNM E11353).

FIGURE 8.—Doraster constellatus, showing the large duck-billed pedicellariae typical of the

Zoroasteridae. Holotype (USNM El 1352).
NUMBER 6 4 11

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FIGURE 9.—Doraster constellatus, dorsal aspect. Holotype (USNM E11352).

FIGURE 10.—Doraster constellatus, ventral aspect. Holotype (USNM El 1352).
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

point of attachment, tapering rapidly to a point; they

may be curved. The 1st or 2nd spine usually bears a

large duckbilled pedicellaria (Figure 8), and there are

numerous small straight pedicellariae. The noncarinate

adambulacrals bear 3 to 5 small spinelets on their outer

surface and do not project into the furrow; they may

also bear small straight pedicellariae.

SIZE RANGE.—R=42 mm to 220 mm; r= 7 mm to

32 mm. Average R/r=10/l. R = major radius, from

center of disc to tip of arm. r=minor radius, from

center of disk to interbrachial margin.

MATERIAL EXAMINED.—Oregon station 3583, 09°

16'N, 81°37'W, 280 fms., May 1962, 1 specimen;

Oregon station 382, 29°11.5'N, 88°07.5'W, 190-210

fms., June 1951,6 specimens; Oregon station 2780, 11°

36'N, 62°52'W, 215-230 fms., April 1960, 2 specimens;

Alaminos station 21/68-A-13, 27°37.5'N, 95°20'W,

350 fms., November 1968, 1 specimen; Oregon II station 10602, 07°46'N, 54°35'W, 299 fms., May 1969,

1 specimen; Albatross station 2376, 29°03'N, 88°16'W,

324 fms., February 1885,8 specimens; Albatross station

2377, 29°07'N, 88°08'W, 210 fms., February 1885, 4

specimens; Albatross station 2396, 28°34'N, 86°48'W,

335 fms., March 1885, 1 specimen; Albatross station

unknown, winter cruise 1885, 1 specimen.

LOCATION OF TYPE-MATERIAL.—Holotype (USNM

El 1352) and 22 paratypes in United States National

Museum; 1 paratype in British Museum; 1 paratype in

Museum of Comparative Zoology at Harvard.

Mammaster sigsbeei (Perrier, 1880)

In view of the fact that some specimens of Doraster

constellatus have previously been misidentified as

Mammaster sigsbeei (Perrier), it may be helpful to

include here a brief redescription of M. sigsbeei.

DESCRIPTION.—The arrangement of plates on the

disc is definite and unmistakable: a centrodorsal plate,

next, 5 interradial plates, then 5 larger radial plates.

In each interradial arc are 2 large somewhat crowdedlooking plates, and over the base of each arm are 3

large tumid plates. All these plates are raised, tumid,

round, somewhat bare (although they may have small

spinelets around the edge). The madreporite is small

and inserted between the 2 interradial arc plates and

the interradial primary plate.

The jaws bear 3 transverse rows of 2, 2, and 3 to 4

long acute spines, and inside the mouth, not visible

without dissection, are 2 more pairs, the central pair

short and blunt, and the outer pair curved away from

the jaws like cow's horns, both pairs covered with small

pedicellariae. Both small and large straight pedicellariae are present in the ambulacral groove and on the

dorsal surface, especially on the disc.

The number of rows of tubefeet diminishes from 4 to

2 about half to three quarters of the way down the arm.

Only the carinal plates and 1 row of plates on each

side extend all the way to the end of the arm. The

terminal plate is as broad as long, with an indentation

FIGURE 11.—Doraster constellatus, dorsal aspect of cleaned specimen. Paratype

(USNM El 1353).
NUMBER 64 13

on the proximal dorsal side. The carinals are overlapped on each side by the plates of the next adjoining

row. These 3 rows of plates are comparatively bare,

with only a few spinules on the distal edges of the

plates, and are probably covered by a thin skin. The

next row of plates is covered with tiny spinules, is overlapped by the adradials, and in turn overlaps the similar row of plates below it. Both of these rows diminish

and disappear before reaching the end of the arm,

the lower row first, then the upper. Next come 2 rows

of narrow elongate plates which disappear distally

before the rows above. Each of the plates in these 4

spinose rows bears 1 larger spine on a turbercle. The

adambulacral plates are alternately carinate and noncarinate, with every other one projecting strongly into

the furrow and bearing 4 stout movable spines in a

transverse row. These spines are frequently curved and

the 1st or 2nd often bears 1 or 2 large pedicellariae.

The adambulacral plates between do not project into

the furrow, and have no furrow spines, but bear 2 to

4 somewhat flattened spines distally side by side in

pairs. The difference between the alternating adambulacrals becomes less distinct distally.

JUVENILE.—A specimen from Combat station 450

has R=9mm. The centrodorsal plate is by far the

largest plate and bears a blunt tubercle. There are

only 2 rows of plates on the arms beyond the carinals

proximally and only 1 row distally. There is no appreciable alteration of the adambulacral plates. There

are only 2 rows of tubefeet the whole length of the

arm. The primary plates of the disc, except for the

centrodorsal plate, are not appreciably raised and

tumid. There are no pedicellariae.

MATERIAL EXAMINED.—MCZ 876, cotype, 1 dry, off

St. Kitts, 208 fms., Blake collection; MCZ Ex. 877,

cotype, 1 dry, 28°42TST, 88°40'W, 321 fms., Blake collection; MCZ 4048, 8 dry, Cuba, Old Bahama Channel off Punta Alegra, 195-230 fms., Atlantis stations

2981B, 2982B, 2982C, March 1938; MCZ 4049, 7 dry,

west end of Old Bahama Channel, 235-260 fms., Atlantis stations 2983, 2983A, March 1938; MCZ 4053,

1 dry, Cuba, off Puerto Tanama, 295 fms., Atlantis

station 3371, April 1938; MCZ 4054, 4 dry, Cuba, Old

Bahama Channel off west end of Cayo Romano, 245-

255 fms., Atlantis stations 3387, 3388, April 1939;

MCZ 4057, 3 dry, Cuba, off Bahia de Matanzas, 285

fms., Atlantis station 3483, May 1939; MCZ 4050, 4

dry, south end of Santoren Channel, southeast of Cay

Sal Bank, 250 fms., Atlantis station 2985; MCZ 4051,

3 dry, Nicholas Channel south of Cay Sal Bank, 280-

300 fms., Atlantis station 2987, March 1938; MCZ

4056, 1 dry, Nicholas Channel south of Cay Sal Bank,

325 fms., Atlantis station 3443, May 1939; MCZ 4055,

13 dry, Cuba, off Caibarien, 245-265 fms., Atlantis

stations 3431, 3434, 3435, 3436, 3437, 3438, April

1939; MCZ 4052, 1 dry, Cuba, off Bahia Matanzas,

170-255 fms., Atlantis station 3000, March 1938;

MCZ 880, cotypes, 2 wet, same as MCZ 876; USNM

E3967, 1 dry, Smithsonian-Johnson expedition station

23, 18°32'N, 66°21'W, 260 fms., 1933; Combat station 450, 10 dry, 23°59'N, 79°43'W, 350 fms., July

1957; Oregon, 5 dry, no station, Florida Keys, 200

fms.; Oregon station 2775, 1 dry, 11°35'N, 62°37/W,

220-230 fms., April 1960.

The Genera and Species of Zoroasteridae

The characters of the other genera of Zoroasteridae

are firmly established by the original authors and by

H. L. Clark (1920) and W. K. Fisher (1919a, b). I

see no need to redescribe any of them here. A list, however, of the genera and species and their known distribution follows.

GENERA AND SPECIES

Zoroaster Thomson, 1873

fulgens Thomson, 1873

actinocles Fisher, 1919

ada mi Koehler, 1909

alfredi Alcock, 1893

angulatus Alcock, 1893

barathri Alcock, 1893

carinatus Alcock, 1893

carinatus philippinensis

Fisher, 1916

evermanni Fisher, 1905

evermanni mordax Fisher,

1919

gilesii Alcock, 1893

hirsutus Ludwig, 1905

longispinus Ludwig, 1905

(?juv.)

macracanthus Clark, 1916

magnificus Ludwig, 1905

microporus Fisher, 1916

ophiactis Fisher, 1916

ophiurus Fisher, 1905

orientalis Hayashi, 1950

perarmatus Clark, 1920

planus Alcock, 1893

DISTRIBUTION

North Atlantic, 120-1600

fms.

Aleutians, 1217 fms.

Bay of Bengal, 569 fms.

Bay of Bengal, 1300-1380

fms.

Indian Ocean, 597-705 fms.

Bay of Bengal, 1520 fms.

Andaman Sea, 130-250 fms.

Borneo, 415 fms.

Southern California, 216—

510 fms.

Washington to California,

275-815 fms.

Andaman Sea, 400-500 fms.

Acapulco, Mexico, 1878 fms.

Eastern Tropical Pacific,

780-1320 fms.

Australia, 250-450 fms.

Gulf of Panama, 1671 fms.

Moluccas, 700 fms.

Celebes, 834 fms.

California, 1059 fms.

Japan, 105-475 fms.

Peru, 536 fms.

Laccadive Sea. 1200 fms.
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

GENERA AND SPECIES

spinulosus Fisher, 1906

fentm Sladen, 1889 (?juv.)

Bythiolophus Fisher, 1916

acanthinus Fisher, 1916

Cnemidaster Sladen, 1889

wyvillei Sladen, 1889

nudus Ludwig, 1905

squameus Alcock, 1893

zea Alcock, 1893

Dor aster, new genus

constellatus, new species

Mammaster Perrier, 1894

sigsbeei (Perrier, 1880)

Myxoderma Fisher, 1905

sacculatus Fisher, 1905

platyacanthus Clark, 1913

platyacanthus rhomaleum

Fisher, 1919

Pholidaster Sladen, 1889

squamatus Sladen, 1889

distinctus Fisher, 1919

Prognaster Perrier, 1891

(genus doubtful)

grimaldii Perrier, 1891

(? seepage 14)

DISTRIBUTION

Hawaii, 328-558 fins.

New Guinea, 1070 fms.

Celebes, 559 fms.

Philippines, 761-1089 fms.

Gulf of California, 998 fms.

Laccadive Sea, 1043 fms.

Indian Ocean, 597-1200 fms.

Caribbean, Gulf of Mexico,

190-350 fms.

Caribbean, 200-450 fms.

West coast of North America,

284-916 fms.

Lower California, 284 fms.

Oregon to California, 223-

550 fms.

Philippines, 100 fms.

Banda Sea, 140 fms.

Azores, 1568 fms.

The Atlantic Species of Zoroaster

In the course of this study, I examined critically the

Atlantic species of the genus Zoroaster. Unfortunately,

the types of many species from other parts of the world

were not available to me, and a complete revision of

the genus was neither possible nor, at this point, particularly necessary. It is appropriate, however, to include here my conclusions regarding the species found

in the Atlantic.

The characters used to distinguish the species of

Zoroaster are, on the whole, unsatisfactory. Degree of

development of the adradial plates is more a function

of growth than a firm taxonomic character; presence

of a central spine on certain plates is untrustworthy,

because of the weakness of attachment (presence or

absence of a tubercle is not necessarily indicative of

presence or absence of a spine) and also because this,

too, may be a function of growth. Large duck-billed

pedicellariae are lacking in Pholidaster and sometimes

in Mammaster, but their number and placement are

variable within the species of Zoroaster. The armature

of the actinolateral plates apparently is also variable

within a species.

As with most asteroids, the number of furrow spines

is a dependable character. Armature of the disc plates,

degree of ensacculate spinulation, and tumidity of disc

plates are mostly useable but less dependable specific

characters. The ratio of minor to major radius may

prove useful and if so, I would be inclined to separate

Perrier's longicaudus from the rest of the Atlantic

Zoroasters. Long slender arms also characterize ackleyi

and seem to set it apart from trispinosus, fulgens, and

diomedeae; however, gradations between ackleyi and

these last three "species" lead one inevitably to the conclusion that R/r is an environment-dependent factor.

Size and placement of papular pores should, perhaps,

be given more attention, but are probably a function

of R/r.

The synonymizing of diomedeae, ackleyi, and trispinosus (and possibly longicaudus) with fulgens has

the unfortunate effect of rendering H.L. Clark's

(1920) key to the species of Zoroaster practically useless. With the types, however, of many of the species

he listed unavailable to me for study and comparison,

I feel it would be unwise to attempt a revised key here.

As this is mostly a deep water family, I am sure that

further synonymizing among the Zoroasteridae, particularly in the genus Zoroaster, is inevitable with further study on a worldwide basis.

H.L. Clark (1920) indicates that Prognaster longicauda Perrier is a Zoroaster; in this he is quite correct.

It is probably Zoroaster fulgens. Neither Clark nor

myself, however, had an opportunity to examine Prognaster grimaldi Perrier, the type species of the genus.

P. grimaldi is known only from the type specimen

(Azores, 1568 fms.), and I suspect, as did Clark, that

it, too, is a Zoroaster, probably fulgens. The only character on which Prognaster is separated from Zoroaster

is that the former has all adambulacral plates alike,

the latter has the usual zoroasterid alternate carinate

and noncarinate adambulacrals. Adambulacral plates

alike would be a very important and unusual character

in this family, but I doubt that it really exists in Perrier's specimen. Perrier's figures in his Monaco report

(1896) show an unusually wide area between the

carinate adambulacrals and the figures look, in every

respect, like Zoroaster fulgens; Perrier, however, states

that he did not observe two kinds of adambulacral

plates. As he had only one specimen, he was probably

reluctant to dissect it sufficiently to determine this

character.
NUMBER 64 15

Zoroaster fulgens Thomson, 1873

Zoroaster ackleyi Perrier, 1883.

Zoroaster diomedeae Verrill, 1884.

Zoroaster trispinosus Koehler, 1895.

Zoroaster bispinosus Koehler, 1909.

DESCRIPTION.—A 5-lobed centrodorsum, 8 primary

radials (in overlapping pairs) and 5 large interradials, plus five large first carinals, make up the disc

dorsum. The anus is between the centrodorsum and 2

primary radials, which are single rather than being

paired like the other 6. The anal pore is surrounded

by small flattened spines. Small pedicellariae may be

scattered over the disc. The madreporite is small, flat,

channeled, and located just distal to a primary interradial.

Each carinal plate on the arms has 2 lobes on each

side and overlaps the carinal plate proximal to it.

Transversely, the carinals overlap the adradials and

the superomarginals overlap both the adradials and

the inferomarginals. Below the inferomarginals are

6-2 rows of actinolateral plates (6 near the disc, becoming 2 by the end of the arm). Each actinolateral

plate bears a long, slender, appressed spine, directed

upward, and mounted on a horseshoe-shaped tubercle,

plus numerous small sacculate spinelets. The inferomarginals are similarly armed. The superomarginals

and carinals frequently (but not always) bear a stout

erect central spine or tubercle, and these plates, as well

as the adradials, are covered with small spinelets. The

carinals are broader than long proximally and longer

than broad distally.

The superambulacral plates are very small and do

not extend beyond about the middle of the arm. The

tubefeet are in 4 rows proximally, becoming 2 rows

less than half way out on the arm. They are stout, conical, terminating in a small suckered disc (except for

those actually within the peristomial cavity, which have

large suckered discs), and the ampullae are double.

The adambulacral plates are alternately carinate and

noncarinate; the carinate adambulacrals bear 5 spines,

2 long slender furrow spines with many small straight

pedicellariae (a large duck-billed pedicellaria may replace the many small ones) and 3 outer spines which

are similar to the 3 small spines of the noncarinate

adambulacrals; these small spines do not bear

pedicellariae.

The jaw, of 4 fused adambulacral plates, bears 2

short stout oral spines on each side in a row along the

oral edge of the jaw, covered with small straight pedicellariae, and behind them are 2 pairs of long acute

spines.

DISCUSSION.—Careful examination of the three socalled species of Atlantic Zoroasters reveals that they

belong to one rather variable species. H. L. Clark

(1920) reviewed the Zoroasteridae very thoroughly

and gave keys to the genera and species. He stated,

quite correctly, that Zoroaster trispinosus Koehler is a

synonym of Z. fulgens; Z. bispinosus Koehler is a

lapsis calami for Z. trispinosus. The other species names

which he listed from the Atlantic as having once been

considered species of Zoroaster have been put into

other genera in the family (Z. sigsbeei=Mammaster

sigsbeei; Z. longicauda=Prognaster longicauda). P.

longicauda, however, is definitely a Zoroaster and must

be returned to that genus. Without more material, it is

not possible to decide if this is a distinct species of

Zoroaster or another synonym of Z. fulgens.

The types of Zoroaster fulgens could not be located;

however, a specimen from Wyville Thomson's own

collection, identified by him and now in the British

Museum (Faroe Channel, 500 fms.) has been examined, as have the types of Z. ackleyi and Z. diomedeae.

In addition, several hundred nontype specimens of the

three so-called species have been examined and compared. Presence, absence, or degree of development of

the superomarginal spines, and relative width of the

carinal plates are the sole characters on which the

Atlantic Zoroasters have been separated. These characters, vague and unsatisfactory to begin with, prove

extremely variable throughout the range of the genus

in Atlantic waters. As the oldest name, Zoroaster fulgens has priority, and Z. ackleyi and Z. diomedeae are

junior synonyms.

On the whole, Zoroaster fulgens from the northern

part of its range tends to be more spinous and more

robust; also, it is generally found below 1000 fathoms.

In the southern part of its range, it is more compact

of skeleton, less spinous, and the arms, because of the

more compact skeleton, seem to be more slender; it is

found in waters as shallow as 200 fathoms.

MATERIAL EXAMINED.—USNM 9066, Albatross station 2052, 39°40'N, 69°21'W, 1098 fms., August 1883,

7 specimens (syntypes of Zoroaster diomedeae) ;

USNM 26258, Albatross station 2043, 39°49'N, 68°-

28'W, 1467 fms., July 1883, 1 specimen (syntype of

Z. diomedeae) ; USNM 14050, Albatross station 2043,

see above, 6 specimens (syntypes of Z. diomedeae);
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

USNM 14049, Albatross station 2095, 39°29'N, 70°-

58'W, 1342 fms., September 1883, 40 specimens (syntypes of Z. diomedeae); USNM 14048, Albatross station 2035, 39°26'N, 7O°O2'W, 1362 fms., July 1883, 73

specimens (syntypes of Z. diomedeae); USNM 9017,

Albatross station 2035, see above, 7 specimens (syntypes

of Z. diomedeae); USNM 9019, Albatross station 2051,

39°41'N, 69°20'W, 1106 fms., August 1883, 5 specimens (syntypes of Z. diomedeae); USNM 9223, Albatross station 2084, 40°16'N, 67°05'W, 1290 fms.,

September 1883, 1 specimen (syntype of Z. diomedeae) ; USNM E4944, Dry Tortugas, Florida, collected

by P. Bartsch, 13 specimens; USNM 14274, Albatross

station 2077, 41'WN, 66°02'W, 1255 fms., September

1883, 1 specimen; USNM 14275, Albatross station

2084, see above, 2 specimens; USNM 7864, Albatross

station 2208, 39°33'N, 7l°16'W, 1178 fms., July 1884,

I specimen; USNM 11265, Martha's Vinyard, Massachusetts, 1390 fms., 2 specimens; USNM 14830, Albatross station 2684, 39^5^ , 70°54'W, 1106 fms., July

1886, 1 specimen; USNM 12041, Albatross station

2571, 40°09'N, 67°09'W, 1356 fms., September 1885,

II specimens; USNM 11263, Albatross station 2562,

S g 0^ ^ , 71°25'W, 1434 fms., August 1885, 54 specimens; USNM 11265, Albatross station 2564, 39^2^ ,

71°23'W, 1390 fms., August 1885, 80 specimens;

USNM 11262, Albatross station unknown, 1885, 10

specimens; USNM 15544, Albatross station 2732,

37°27'N, 73°33'W, 1152 fms., October 1886, 6 specimens; USNM 17999, Albatross station 2196, 39°35>N,

69°44'W, 1230 fms., August 1884, 1 specimen; all of

the above specimens were identified by A. E. Verrill

as Zoroaster diomedeae.

USNM E5696, South of Dry Tortugas, Florida, collected by W. Schmitt, August 1932, 1 specimen;

USNM 18507, Albatross station 2655, 27°22'N, 78°-

07'W, 338 fms., May 1886, 5 specimens; USNM

10422, Albatross station 2394, 28°38'N, 87°02'W, 420

fms., March 1885, 2 specimens; USNM 18450, Albatross station 2396, 28°34'N, 86°48'W, 335 fms., March

1885, 2 specimens; USNM 18504, Albatross station

2659, 28°32'N, 78°42'W, 509 fms. May 1886, 1 specimen; USNM 18505, Albatross station 2380, 28°-

02'N, 87°43'W, 1430 fms., March 1885, 1 specimen;

USNM 18506, Albatross station 2381, 28°05/N, 87°-

56'W, 1330 fms., March 1885, 1 specimen; USNM

38243, Fish Hawk station 7281, 24°13'N, 81°58'W,

304 fms., February 1902, 1 specimen; Alaminos station 3C/68-A-7, 27°42'N, 87°44/W, 1500 fms., July

1968, 1 specimen; Oregon station 1537, 24°29'N,

83°27'W, 212 fms, June 1956, 2 specimens; Oregon

station 2574, 26°34'N, 89°53'W, 1450 fms, July 1959,

4 specimens; Oregon station 3561, 16°35'N, 80<WW,

400 fms. May 1962, 1 specimen; Oregon station 3560,

16°35TSr, 80°10'W, 315 fms. May 1962, 2 specimens;

Oregon station 5929, 15°39'N, 61°10'W, 335 fms,

March 1966, 5 specimens; Oregon station 4413, 11°-

53'N, 69°25'W, 350 fms, October 1963, 3 specimens;

Oregon station 2571, 26°30'N, 9O°31'W, 1300 fms,

July 1969, 1 specimen; Oregon station 2202, 28°58'N,

88°11'W, 625 fms, June 1958, 2 specimens; Oregon

station 2779, 11°35'N, 62°59'W, 260 fms, April 1960,

1 specimen; Oregon station 2824, 29°75'N, 88°04'W,

365 fms, July 1960, 2 specimens; Oregon station

2652, 18°18'N, 67°18'W, 300 fms, October 1959, 10

specimens; Oregon station 1538, 24°29'N, 83°32'W,

200 fms, June 1956, 6 specimens; Oregon station 382,

29°11'N, 88°07'W, 200 fms, June 1951, 6 specimens;

Oregon station 2650, 18°16'N, 67°17'W, 250 fms,

October 1959, 5 specimens; all of the above specimens

were identified by various people as Zoroaster ackleyi.

British Museum (Natural History) 1949-2-14-29,

Faroe Channel, 500 fms, 1 specimen, collected by

Wyville Thomson; USNM 33308, west coast of Ireland,

gift of Department of Agriculture, Ireland, 2 specimens; USNM 7865, Albatross station 2206, 39°35'N,

71°24'W, 1043 fms, August 1884, 1 specimen; USNM

31265, Fish Hawk station unknown, San Juan, Puerto

Rico, 1 specimen; USNM 31266, same as 31265, 1

specimen; Alaminos station 1/68-A-13, 25°42'N,

96°04'W, 480 fms, November 1968, 2 specimens;

Alaminos station 10B/68-A-3, 25°12'N, 96°21'W, 550

fms, March 1968, 2 specimens; Oregon station 3654,

29°08'N, 88°00'W, 400 fms, July 1962, 6 specimens;

Oregon II station 10619, O7°51'N, 54°23'W, 425 fms.

May 1969, 1 specimen.

MCZ 873, Santa Cruz, 248 fms, Blake collection, 4

specimens (syntypes of Z. ackleyi) ; MCZ 4045, Cuba,

off Bahia de Matanzas, 145-255 fms, Atlantis stations

2999, 3000, March 1938, 19 specimens; MCZ 4047,

Cuba, off Bahia de Matanzas, 175-285 fms, Atlantis

stations 3465, 3467, 3478, 3479, 3480, 3482, 3483, May

1939, 23 specimens; MCZ 4041, Cuba, Old Bahama

Channel, 105-260 fms, Atlantis stations 2980A, 2982B,

2982C, March 1938, 5 specimens; MCZ 4044, Cuba,

Nicholas Channel off Bahia de Santa Clara, 475-580

fms, Atlantis stations 2991A, 2992A, 2993, March

1938, 3 specimens; MCZ 874, off Yucatan, 600 fms.
NUMBER 6 4 17

Blake collection, 1 specimen; MCZ 3046, South Tortugas, 240 fms., 5 specimens; MCZ 4046, Cuba, off

Caibarien, 260 fms., Atlantis station 3437, May 1939,

2 specimens; MCZ 4040, Cuba, off Punta Lucrecia,

1015 fms., Atlantis station 2975, March 1938, 1 specimen; MCZ 4042, Cuba, west end of Old Bahama

Channel off north coast, 235-260 fms., Atlantis station

2983A, March 1938, 2 specimens; MCZ 4043, Nicholas

Channel, south of Cay Sal Bank, 280-300 fms., Atlantis

station 2987, March 1938, 1 specimen; MCZ 879,

Santa Cruz, same as 873, 3 specimens (syntypes of

Z. ackleyi); MCZ 4075, same as 4047, 6 specimens;

all of the above were identified as Zoroaster ackleyi.

MCZ 3719, off Nantucket, 40°05'N, 67°52'W, 1325

fms., Atlantis station unknown, 1939, 37 specimens;

MCZ 3744, 40°06'N, 68°06'W, Atlantis, July 1939,

1160-1100 fms., 8 specimens; MCZ 872, Albatross

station 2043, Gulf Stream, 4 specimens; MCZ 878,

south of Nova Scotia, Challenger collection, 1 specimen; MCZ 949, 38°96'N, 73°10'W, 1186 fms., Blake

collection, 2 specimens; all of the above specimens

were identified as Zoroaster diomedeae. MCZ 2730,

northwest of Azores, 1000 fms., 1903, 2 specimens,

identified as Zoroaster trispinosus.

MCZ 1856, North Atlantic, Thor collection, 1906,

1 specimen. I also examined MCZ 875, Eastern

Atlantic, Talisman collection, 1 specimen, which was

identified by Perrier (1894) as Prognaster longicauda;

as H. L. Clark suggests, it is definitely a Zoroaster,

but because of the disportionately small disc and long

slender arms, I hesitate to synonymize it with Z. fulgens.

Acknowledgments

I am especially grateful to Mr. Jerald Halpern of the

University of Miami for his valuable comments and

suggestions during the preparation of this paper. For

the opportunity to examine material in collections

other than those of the United States National

Museum, I wish to thank Miss Ailsa M. Clark, of the

British Museum (Natural History); Professor H. B.

Fell, of the Museum of Comparative Zoology at

Harvard; and Dr. A. N. Soloviev, of the Paleontological Institute in Moscow, who kindly prepared and sent

me a plaster cast of part of the type of Calliasterella

mira. I am indebted to Dr. Klaus Ruetzler, of the

United States National Museum, for the photographs,

and to Dr. David L. Pawson, Division of Echinoderms,

and Dr. Porter M. Kier, Department of Paleobiology, for reading and most helpfully criticizing the

manuscript.

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18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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