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Vol te 3 October 1060

PROCEEDINGS
OF THE

) i
]

BIOLOGICAL SOCIETY OF WASHINGTON,

FIVE NEW PARASITIC COPEPODS FROM
CALIFORNIA INSHORE FISH

By RoceEr F. CRESSEY
Smithsonian Institution, Washington, D. C.

A collection of parasitic copepods sent to me by Mr. Ed-
mund Hobson of the U. S. Bureau of Sport Fisheries and Wild-
life, U.S. Department of Interior, contained a number of new
species. One of these (Caligus hobsoni Cressey) has been re-
ported elsewhere. This paper reports on five more, one of
which constitutes a new genus of taeniacanthid. The entire col-
lection plus additional new species will be reported on later.

All material has been deposited in the Smithsonian Institu-
tion, Division of Crustacea.

Mr. Hobson was assisted in the collecting by Mr. Lloyd D.
Richards of the U. S. Bureau of Sport Fisheries and Wildlife.

Taeniastrotos new genus

Diagnosis: Bomolochiidae, Taeniacanthinae. First thoracic segment
fused to head. Thoracic segments 2, 3, and 4 free. Postantennal process
(maxillary hook) present. First antenna 7-segmented. Padlike process
present between bases of first antenna on ventral rostral area. Second an-
tenna, mandible, and first maxilla as in other members of the subfamily.
Second maxilla with reduced terminal segment bearing 3 weak setae. Max-
illiped well-developed and posterior to mouthparts. Rami of legs 1-4 3-
segmented (segmentation of the rami of leg 1 incomplete). Terminal en-
dopod segment of leg 4 without lateral spine. Leg 5 2-segmented, armed
as in other members of the family. Leg 6 represented by 3 setae on the
area of egg sac attachment.

The male is unknown.

Type species: Taeniastrotos californiensis new speices.

Etymology: From Greek, masc. The generic name is a combination of
Taeniacanthus and Anchistrotos, the two genera to which the new genus
seems most closely related.

31—Proc. BioL. Soc. WaAsH., VoL. 82, 1969 (409 )

410 Proceedings of the Biological Society of Washington

Remarks: The new genus can be separated from all other genera of the
Taeniacanthinae by the reduced nature of the second maxilla and the lack
of a lateral spine on the last endopod segment of leg 4. This is the first
record of a taeniacanthin from the eastern Pacific and it is not surprising
that it would be an undescribed form when one considers the high rate of
endemism in that area.

Taeniastrotos californiensis new species

Figures 1-11

Material studied: Holotype @ (USNM 126240), and 7 paratype @ ?
(USNM 126241) were collected from the body surface of Paralebrax
nebulifer (Giard) at La Jolla, California 2 October 1968.

Female: Body form as in figure 1. Total length 1.5 mm. Greatest width
0.65 mm (measured at widest part of cephalon). Cephalon comprises a
little more than one-third total body length. First thoracic segment com-
pletely fused with head. Thoracic segments bearing legs 2, 3, and 4 free;
each narrower than the one before. Genital segment small, only slightly
longer than first abdominal segment and somewhat wider than long. Ab-
domen 4-segmented, 89 u, 83 uw, 59 uw, and 83 w in length respectively.
Caudal rami (fig. 2) about three times as long as wide (94 u x 30 4);
armed with one lateral, four terminal, and one inner subterminal setae,
longest seta 519 uw in length.

First antenna (fig. 3) 7-segmented; armed with setae as in the figure,
an aesthete present on the penultimate segment. A padlike process, some-
what triangular, its broadest portion projecting slightly beyond the ante-
rior margin of the body, present between the bases of the first antennae on
the ventral rostral area (see fig. 3). A heavily sclerotized postantennal
process present (see fig. 3). Second antenna (fig. 4) similar to that of
other species in the subfamily; bearing one short thick spine, three longer
spines, and three setae at tip (longest bearing a few plumosities ).

Mandible (see fig. 5) a simple bladelike appendage bearing two un-
equal broad spines at tip. First maxilla (see fig. 5) consisting of a small
lobe bearing a long plumose seta directed posteriorly, a bladelike blunt
seta directed anteriorly, and two smaller setae. Second maxilla (see fig.
5) 2-segmented; terminal segment small, bearing three weak setae. Max-
illiped (fig. 6) basal segment unarmed, terminal segment in form of a
stout claw.

Legs 1-4 biramose. Leg 1 (fig. 7) exopod with plumose setae on the
outer distal corners of each of the first two segments; terminal segment
with two weak and five well-developed setae: endopod segmentation in-

>

Fics. 1-6. Taeniastrotos californiensis n. gen., n. sp. female: 1, dorsal;
2, caudal ramus, ventral; 3, first antenna; 4, second antenna; 5, mouth-
parts; 6, maxilliped.

41]

New California parasitic copepods

—

=

412 Proceedings of the Biological Society of Washington

complete but ramus bearing eight well-developed setae. Leg 2 (fig. 8)
exopod spines on outer distal corners of first two segments plus first spine
on last segment with conspicuously serrate edges; endopod second segment
with spinelike process on outer distal corner, last segment distalmost spine
finely plumose and sclerotized on basal half but distal half hyaline and
naked. Basipod of leg 2 with two comblike rows of spines. Leg 3 (fig. 9)
similar to leg 2 but with only one row of comblike spines on basipod, and
additional serrate spine on last exopod segment, and terminal spine on last
exopod segment normal. Leg 4 (fig. 10) exopod with serrate spines on all
three segments, otherwise armed as in the figure; endopod last segment
with a short terminal spine on outer distal corner and a long seta on inner
distal corner, no lateral spine. Leg 5 (fig. 11) 2-segmented; basal segment
with a long plumose seta and a row of spinules on outer distal corner; last
segment with an outer lateral spine and an inner and outer spine ter-
minally separated by a slightly longer seta, a small patch of spinules is
present near the base of the inner spine. Leg 6 represented by two long
setae on the midlateral margins of the genital segment at the area of egg
sac attachment.

Egg sacs long (1.1 mm) and containing about 100 eggs arranged in
two rows.

Male: Unknown.

Remarks: The species is named for the type locality (California). An-
chistrotos pleuronichthydis Yamaguti 1939 has a first maxilla with the
blunt anteriorly directed seta as in the new species described here. How-
ever, it differs from californiensis on other points and may represent a form
intermediate between the new genus described here an Anchistrotos.

Bomolochus longicaudus new species
Figures 12-22

Material studied: Holotype 2 (USNM 126242), a single @ paratype
(USNM 126243) was collected from the gill cavity of Paralebrax nebulifer
(Giard) 8 August 1968, and additional material from the same host (1 @
6 August 1968, 3 9 2 1 October 1968, and 1 @ 2 October 1968). An
additional female was collected from the gill cavity of Paralebrax clathra-
tus (Giard) 10 October 1968. All material was collected at La Jolla, Cali-
fornia.

>

Fics. 7-11. Taeniastrotos californiensis n. gen., n. sp., female (cont.):
7, first leg; 8, second leg; 9, third leg; 10, fourth leg; 11, fifth leg.

Fics. 12-16. Bomolochus longicaudus n. sp., female: 12, dorsal; 13,
caudal rami, ventral; 14, first antenna; 15, second antenna; 16, mouth-
parts.

Fics. 17-20. Bomolochus longicaudus n. sp., female (cont.): 17, maxil-
liped; 18, first leg; 19, second leg; 20, third leg.

New California parasitic

copepods

414 Proceedings of the Biological Society of Washington

it

———

415

New California parasitic copepods

416 Proceedings of the Biological Society of Washington

Female: Body form as in figure 12. Total length 2.03 mm. Greatest
width 1.01 mm (measured at widest part of cephalon). Cephalon com-
prises about one-third total body length. First thoracic segment fused with
head. Thoracic segments bearing legs 2-4 free. Genital segment about as
wide as long (approx. 200 1). Abdomen 3-segmented; 147 u, 136 uw, and
124 uw in length respectively, second and third segments each slightly nar-
rower than preceding one, last segment with two patches of spinules ven-
trally. Caudal rami (fig. 13) about three times as long as wide (162 w x
50 «); each ramus with four terminal, one subterminal, and one lateral
setae, longest seta 600 u long.

First antenna (fig. 14) with five distinct segments, second segment
weakly subdivided making the actual number of segments difficult to as-
certain; each segment armed with setae as in the figure. Second antenna
(fig. 15) basically as in other members of the genus, spinules on surface
not arranged in definite rows. Mouthparts as in other members of the
genus except first maxilla (see fig. 16); first maxilla with very long inner
seta, reaching nearly to the base of first legs. Maxilliped (fig. 17) terminal
claw with accessory spine.

Legs 1-4 biramose. Leg 1 (fig. 18) exopod segmentation incomplete,
two weak spines present on outer margin of ramus and six terminal to in-
ner setae; endopod 3-segmented, first two segments with an inner seta on
each and last segment with five terminal setae. Leg 2 (fig. 19) exopod
outer five spines on ramus without fringes or hairs, terminal spine with
fine hairs along inner margin; exopod last segment with two short outer
spines each with a hyaline fringe, both rami with additional setae as in
the figure. Leg 3. (fig. 20) similar to leg 2 except for a reduction in the
number of spines and setae. Leg 4 (fig. 21) exopod similar to legs 2 and 3;
endopod first two segments each with a short seta on inner distal corner,
last segment with an inner and an outer short spine and a short terminal
seta between. Leg 5 (fig. 22) 2-segmented; basal segment wtih a patch
of spatulate spinules on distal third, last segment armed as in other mem-
bers of the genus except for patches of spatulate spinules as shown in the
figure. Leg 6 represented by three long setae at the point of egg sac at-
tachment.

None of the material was ovigerous.

Male: Unknown.

Remarks: This species may be separated from all other species of the
genus, except the following new species, by the nature of the first maxilla
with its exceptionally long inner seta. B. longicaudus can be easily sep-
arated from the following new species on the basis of characters to be dis-
cussed following the description of this second species of Bomolochus.

=

Fics. 21-25. Bomolochus longicaudus n. sp., female (cont.): 21, fourth
leg; 22, fifth leg. Bomolochus prolixus n. sp., female: 23, dorsal; 24,
caudal ramus, ventral; 25, first antenna.

417

New California parasitic copepods

418 Proceedings of the Biological Society of Washington

Bomolochus prolixus new species
Figures 23-32

Material studied: Holotype @ (USNM 126244), 4 paratype 9? 9
(USNM 126245) were collected from the gill cavity of Pleuronichthys
coenosus Giard, and additional material from the same host (1 @ 12 July
1968, 1 @ 6 August 1968, 2 ? 2 6 August 1968, 1 @ 16 October 1968,
and 1 @ 15 January 1969). All material was collected at La Jolla, Cali-
fornia.

Female: Body form as in figure 23. Total length 1.6 mm. Greatest
width 0.58 mm (measured at widest part of cephalon). Cephalon com-
prises about one-fourth total body length. First thoracic segment fused
with head. Thoracic segments bearing legs 2—4 free. Genital segment
wider than long (132 uw X 103 «). Abdomen 3-segmented; 118 y, 94 uy,
and 118 « in length respectively, second and third segments each slightly
narrower than preceding one. Caudal rami (fig. 24) longer than wide
(100 » xX 65 «); bearing setae as in the previous species and two rows of
spinules, spinules of the outer row of the usual pointed nature but the
inner row spinules are spatulate (similar rows occur on the last abdominal
segment ).

First antenna (fig. 25) similar to B. longicaudus, second, third and
fourth segments weakly articulated giving the appearance of a long sec-
ond segment. Second antenna (fig. 26) also similar to preceding species
but spinules on surface arranged in definite rows. Mouthparts as in B.
longicaudus. Maxilliped (fig. 27) terminal hook with accessory process.

Legs 1-4 biramose. Leg 1 (fig. 28) exopod 3-segmented and clearly
articulated; outer spine on first segment somewhat recurved with heavy
fringe on outer edge and attenuated at tip, spine on second segment and
first two spines on last segment with sclerotized basal half, a hyaline distal
portion and a small discrete tip giving whole spine the appearance of
being composed of three parts: endopod as in B. longicaudus. Leg 2 (fig.
29) as in B. longicaudus except terminalmost spines on exopod with an
outer hyaline fringe. Leg 3 (fig. 30) as in B. longicaudus except for patch
of spinules on basipod and other minor variations in armature as indicated
by the figure. Leg 4 (fig. 31) armed as in B. longicaudus but endopod
segments not elongated as in longicaudus. Leg 5 (fig. 32) 2-segmented;
first segment with large patch of spinules on outer distal corner, second
segment with usual lateral seta and three terminal spines or setae, inner
and outer spines pinched at distal third, terminal and outer borders with
spinules as in the figure. Leg 6 represented by three long setae at area of
egg sac attachment.

None of the material was ovigerous.

Male: Unknown.

>

Fics. 26-30. Bomolochus prolixus n. sp., female (cont.): 26, second

antenna; 27, maxilliped; 28, exopod of first leg; 29, second leg; 30, third
leg.

420 Proceedings of the Biological Society of Washington

Remarks: This species, like B. longicaudus, can be separated from all
other species of the genus on the basis of the first maxilla. It can be sep-
arated from longicaudus by the nature of the caudal rami, the distinct rows
of spinules on the second antenna, the nature of the fourth leg endopod,
and the many other variations in the armature of the legs.

Bomolochus spinulus new species
Figures 33-45

Material studied: Holotype 2 (USNM 126246) and 43 paratype 2 9
(126247) from the gili cavity of Scorpaena guttata Giard collected at La
Jolla, California, 10 October 1968. Other material from the same host (2
2 12 July 1968, 1 @ 14 October 1968, 39 2 9 15 October 1968, 5 2 9
16 October 1968, and 29 2 2 16 October 1968) from Oxylebius pictus
Gill (1 @ 9 December 1968) from Sebastodes mystinus (Jordan and Gil-
bert) (3 9 @ 24 September 1968 and 1 ¢ 14 October 1968) and from
Sebastodes serranoides Eigenmann and Eigenmann (8 2 2 12 September
1968 and 1 @ 12 September 1968). All other material collected at La
Jolla, California.

Female: Body form as in figure 33. Total length 1.49 mm. Greatest
width 0.87 mm (measured at widest part of cephalon). Cephalon slightly
less than one-third body length. Thoracic segments bearing legs 24 free.
Thoracic segment bearing leg 4 small and scarcely visible dorsally. Geni-
tal segment about as wide as long (200 4). Abdomen 3-segmented, 224
u, 200 w, and 112 uw long respectively. Caudal rami about twice as long as
wide (142 u x 60 ~); longest seta 768 u long, patch of spinules on ventral
distal surface (comparable patch on last abdominal segment also).

First antenna (fig. 35) armed as in the figure, an aesthete present on
penultimate segment; segmentation weak giving the appearance of 5 seg-
ments. Second antenna (fig. 36) generally as in other members of the
genus except claw at base of last segment well-developed and in the form
of a hook. Mandible (fig. 37) bladelike with two fringed processes at tip.
Labrum with two lateral patches of spinules on posterior corners. First
maxilla of usual form with two inner long setae and two outer short ones.

>

Fics. 31-37. Bomolochus prolixus n. sp., female (cont.): 31, fourth leg;
32, fifth leg. Bomolochus spinulus n. sp., female: 33, dorsal; 34, caudal
ramus; 35, first antenna; 36, second antenna; 37, mandible.

Fics. 38-44. Bomolochus spinulus n. sp., female (cont.): 38, parag-
nath; 39, first maxilla; 40, maxilliped hook; 41, first leg; 42, second leg;
43, third leg; 44, fourth leg.

Fics. 45-52. Bomolochus spinulus n. sp., female (cont.): 45, fifth leg.
Lepeophtheirus paulus n. sp., female: 46, dorsal; 47, abdomen and caudal
rami, ventral; 48, first antenna; 49, postantennal process; 50, second an-
tenna; 51, postoral process; 52, maxilliped.

New California parasitic copepods

Zz
ZB

421

422 Proceedings of the Biological Society of Washington

423

New California parasitic copepods

424 Proceedings of the Biological Society of Washington

Paragnath in the form of a lobe with a patch of spinules and a striated tip
(fig. 38). First maxilla (fig. 39) as in other members of the genus. Maxil-
liped hook (fig. 40) S-shaped and with an accessory process.

Legs 14 biramose. Leg 1 (fig. 41) both rami clearly 3-segmented,
rami armed as in the figure. Leg 2 (fig. 42) exopod last segment two ter-
minalmost spines with spinose fringe, penultimate spine somewhat over-
lapping ultimate spine: otherwise armed in the usual manner. Leg 3 (fig.
43) exopod spines on first two segments angular in shape and recurved in-
wardly, first two spines on last segment similar in nature but not as con-
spicuously recurved. Leg 4 (fig. 44) similar to other members of the
genus except for prominent patches of spinules on outer distal corners of
endopod segments. Leg 5 (fig. 45) 2-segmented and armed in usual
manner, patches of spinules on both segments as in the figure. Leg 6 rep-
resented by three setae at the area of egg sac attachment.

Egg sac containing approximately 100 eggs and about 0.95 mm long.

Male: Unknown.

Remarks: This new species can be distinguished from all other species
of Bomolochus except B. attenuatus Wilson 1913 by the nature and posi-
tion of the outer spines on the exopod of leg 3. Bomolochus spinulus is
closely related to B. attenuatus. The nature of the outer spines on leg 3 is
the same in both species. Both species have patches of spinules on the en-
dopod of leg 4 plus a patch of spinules on the ventral posterior surface of
the caudal rami. They can be separated based on the following differ-
ences: the hooklike spine on the second antenna of attenuatus extends to
the tip of the terminal setae whereas that of spinulus is only half the length,
the last abdominal segment of attenuatus bears a row of spinules on the
posterior ventral surface as opposed to a patch of spinules in the same
area on spinulus. The holotype of B. attenuatus was examined for this com-
parison. Bomolochus attenuatus was described from Scorpaena plumieri
from Jamaica.

Lepeophtheirus paulus new species
Figures 46-61

Material studied: Holotype 2 (USNM 126248), allotype @ (USNM
126249), and 4 paratypes (USNM 126250, 3 22,1 &) were collected
from the roof of the mouth of Sebastodes serriceps (Jordan and Gilbert)
at La Jolla, California 22 July 1968. Additional material was collected
from the same species of host and at the same locality (1 9,4 292 164,
1 ? 19 July 1968; 2 99 1 $ 22 July 1968; 2 99,3 22 26 July 1968;
20 2 2 30 September 1968; 6 2 2 10 October 1968).

>

Fics. 53-58. Lepeophtheirus paulus n. sp., female (cont.): 53, sternal
furca; 54, first leg; 55, second leg; 56, third leg; 57, fourth leg; 58, fifth
leg.

New California parasitic copepods 425

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426 Proceedings of the Biological Society of Washington

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Fics. 59-61. Lepeophtheirus paulus n. sp., male: 59, dorsal; 60, genital
segment, abdomen, and caudal rami; 61, second antenna.

Female: Body form as in figure 46. Total length 1.3 mm. Greatest
width 1.3 mm (measured at widest part of cephalon). Five specimens
measured for length and width all measured the same as above. Cephalon
comprising about one-half total body length. Genital segment nearly
round, slightly wider than long (1.2 x 1.0 mm). Abdomen (see fig. 47)
short (159 «) and 1-segmented. Caudal rami (see fig. 47) short and about
as wide as long, bearing one lateral, 2 subterminal, and three terminal
plumose setae, longest seta 336 u long.

First antenna (fig. 48) 2-segmented and armed as in the figure. Post-
antennal process (fig. 49) with single outer hooklike process, 2 sensory
hairs on base and a sensory hair with terminus split into four hairs lo-
cated near base. Second antenna (fig. 50) with well-developed hook
bearing two setae. Mandible as in other members of genus bearing 12
teeth. First maxilla (fig. 51) a simple lobe bearing three setae. Second
maxilla as in other members of the genus. Postoral process (see fig. 51)
bifurcate, outer tine larger than inner. Maxilliped (fig. 52) hooklike, a
rugose area present on surface of basal segment. Sternal furca (fig. 53)
with rounded tines.

Legs 1-3 biramose. Leg 1 (fig. 54) exopod with three terminal spines
and four terminal to inner setae, endopod a simple, weakly developed proc-
ess near base of exopod. Leg 2 (fig. 55) rami 3-segmented; exopod seg-
ments with a total of four well-developed spines on outer margins and
spines oriented along same axis as ramus, endopod with setae as in the
figure. Leg 3 (fig. 56) exopod basal segment with a well-developed spine
oriented so as to lie over the following segment, endopod 2-segmented and
both rami armed with setae as in the figure. Leg 4 (fig. 57) last segment
with three terminal setae, median seta longest. Leg 5 (fig. 58) located at

New California parasitic copepods 427

posterior corners of the genital segment and consisting of an anterior lobe
bearing a single seta and a posterior lobe bearing three setae of about
equal length. Leg 6 absent.

Egg sac short (1.45 mm) and bearing about 12-15 eggs each.

Male: Body form as in figure 59. Total length and width of three spec-
imens 1.4 x 0.85 mm, 1.5 x 0.98 mm, and 1.4 x 0.82 mm. Cephalon
comprising about three-fourths total body length. Genital segment wider
than long (442 u x 384 uw). Abdomen (fig. 60) 1-segmented and short
(106 «). Caudal rami armed as in the female. Appendages as in the
female except second antenna (fig. 61) with rugose patches on basal
segments and accessory tooth on claw. Leg 6 represented by three plu-
mose setae near junction of genital segment and abdomen.

Remarks: This new species can be separated from all other species of
Lepeophtheirus by the nature of the fourth leg. In L. paulus the middle
seta is longer than either of the other two. Typically the outer seta is
longest or at least as long as the middle. In other respects this species is
most closely related to L. elegans Gussev 1951.

LITERATURE CITED

Cressey, R. F. 1969. Caligus hobsoni, a new species of parasitic copepod
from California. Jour. Parasitology, 55(2): 431-434.

Gussev, A. B. 1951. Parasitic Copepoda of some marine fishes. Collected
Papers on Parasitology from the Zoological Institute, Academy
of Science SSSR, XIII: 394-463.

Witson, C. B. 1913. Crustacean parasites of West Indian fishes and land
crabs, with descriptions of new genera and species. Proc. U.S.
Nat. Mus., 44 (1950): 189-277.

YamacutTl, S. 1939. Parasitic copepods from fishes of Japan Part 4. Cy-
clopoida, II. Volumen Jubilare Pro. Prof. S. Yoshida, Vol. II:
391-415.

428 Proceedings of the Biological Society of Washington

nil 38 3 October 1969 _

PROCEEDINGS —
OF THE ( OCI ZC
BIOLOGICAL SOCIETY OF WASHINGTON\ (igaarii2~

THE GENERA STHENELANELLA MOORE AND
EULEANIRA HORST (POLYCHAETA, SIGALIONIDAE )

By Marran H. PETTIBONE
Smithsonian Institution, Washington, D.C.

In connection with a review of the genera in the polychaete
family Sigalionidae, it was determined that Euleanira Horst
(1916) should be referred to Sthenelanella Moore (1910).
Four species have heretofore been referred to the two genera.
They include the following:

Sthenelanella Moore, 1910: S. uniformis Moore, 1910. Cali-
fornia.

S. atypica Berkeley and Berk-
eley, 1941. Southern Cali-
fornia. Referred to S. wuni-
formis (see below).

S. polymorpha Hartmann-
Schroder, 1962. Chile. Not
Sthenelanella (see page 437).

Euleanira Horst, 1916: £. ehlersi Horst, 1916. Dutch
East Indies. Referred to
Sthenelanella (see below).

Type-specimens of S. wniformis and S. atypica, deposited in the
Smithsonian Institution (USNM), E. ehlersi, deposited in the
Zoological Museum Amsterdam (ZMA) and Rijksmuseum Nat-
ural History Leiden (RNHL), were re-examined.

This study was aided in part by a grant from the National
Science Foundation (GB-1269). For the loan of type-speci-
mens, I wish to extend thanks to S. van der Spoel of the Zoo-
logical Museum Amsterdam and to J. van der Land of the
Rijksmuseum Natural History Leiden. The manuscript bene-
fited from the suggestions of M. L. Jones and J. L. Barnard,
both of the Smithsonian Institution.

32—Proc. Biot. Soc. WaAsH., VoL. 82, 1969 (429 )

430 Proceedings of the Biological Society of Washington

6 ————
DP ee SS
oe SS

g

Fic. 1. Sthenelanella uniformis (Syntypes of S. atypica, USNM 32849):
a, Anterior end, dorsal view, elytra on segments II and IV removed;
pharynx partially extended causing tentacular parapodia to be spread
apart; posterior part of prostomium hidden by segment II; b, anterior end,
ventral view; c, prostomium and tentacular parapodium, lateral view; d,
tentacular parapodium, inner view, showing tentacular lamella; e, para-
podium from segment II, posterior view; f, neurosetae from same; g, para-
podium from segment III, anterior view. (au, auricle; br, branchia; ct,
ctenidia; ] An, lateral antenna; t La, tentacular lamella).

=

0.5mm

Genera Sthenelanella and Euleanira 431

FAMILY SIGALIONIDAE
Genus Sthenelanella Moore, 1910

Type-species: S. uniformis Moore, 1910, by monotypy. Gender: femi-
nine Synonym: Euleanira Horst, 1916. Type-species: E. ehlersi Horst,
1916, by monotypy. Gender: feminine.

Diagnosis: Body slender, depressed, with segments up to 75. Elytra
numerous pairs, arranged on segments 2, 4, 5, 7, then alternate segments to
25 or 27, then continuing on all segments to end of body. Branchiae short,
conical, lateral to dorsal tubercles or elytrophores from segment II on.
Without dorsal cirri. Prostomium rounded, fused with tentacular segment
(1); ceratophore of median antenna with lateral auricles and short style;
lateral antennae very short, fused to tentacular parapodia; palps long,
slender, tapered, emerging lateral to tentacular parapodia, without palpal
sheaths; 2 pairs eyes on raised ocular areas lateral to ceratophore of me-
dian antenna; tentacular parapodia uniramous, extending anteriorly medial
to palps, each with single aciculum, 2 tentacular cirri, well-developed fan-
shaped bundles of capillary setae, and medial tentacular lamella. Parapo-
dia of segments II-IV directed anteriorly; buccal segment (II) with ven-
tral buccal cirri longer than following ventral cirri. Parapodia biramous,
with rami closely united, with notopodial ctenidia; without parapodial
stylodes. Notopodia with conical acicular lobes and inflated rounded
upper lobes; notosetae simple, capillary, spinous; beginning about segment
16, additional long threadlike notosetae formed from notopodial spinning
glands. Neuropodia with rounded presetal and postsetal lobes. Neuro-
setae forming vertical bundles, all compound, with blades short, sickle-
and rod-shaped; blades of neuropodia II-IV longer; some with spinous
stems. Ventral cirri slender, tapered and subulate. Pharynx with 13 or
more pairs distal papillae and 2 pairs interlocking teeth. Occupy long,
tough, fibrous tubes.

Sthenelanella uniformis Moore
Figures 1-3

Sthenelanella uniformis Moore, 1910, p. 391, pl. 33, figs. 105-112.—
Hartman, 1939, p. 69, pl. 18, figs. 226-231; 1961, p. 54; 1968, p. 169,
figs. 1-6.

Sthenelanella atypica Berkeley and Berkeley, 1941, p. 26, pl. 5, figs. 1-3.

Stenelanella [sic] uniformis.—Reish, 1968, p. 72.

Material examined: CALIFORNIA, Albatross in 1904 (exact locality
unknown )—Holotype S. uniformis (USNM 17385). SOUTHERN CALI-
FORNIA, G. E. MacGinitie, collector, off Corona del Mar, 22—31 meters;
off Balboa—34 Syntypes S. atypica (USNM 3248-32850 ).

Description: As in generic diagnosis. Length up to 26 mm, width 2-3
mm, including parapodia, and 3-4 mm, including setae; segments up to
70. Elytra delicate, transparent, on all segments from segment 27 on; first
elytral pair round, with fringe of short, crowded papillae on anterior mar-
gin; rest of elytra subreniform to oval, smooth or with scattered sensory

432 Proceedings of the Biological Society of Washington

O.lmm

NN,

= E
=
™ ive)
3 (} 4 fa
wees = ey gi dt aed hy Y
—— = YEE:
=— = IZLE =
== — _ FE
aa --25-+ (WS
s VSS
= 2 OQ#$—$
\ Se ee 7 =TWAWAS
SS x

Fic. 2. Sthenelanella uniformis (Syntypes of S. atypica, USNM 32849):
a, First elytron; b, anterior elytron; c, parapodium from segment IV, pos-
terior view; d, neurosetae from same; e, parapodium from anterior region
(segment 13), anterior view; f, neurosetae from same.

papillae on lateral margin; anterior elytra with rust-colored mottled pig-
mentation (fig. 2a, b). Prostomium with posterior part hidden dorsally by
segment II, with ceratophore of median antenna equipped with lateral
auricles in middle of ceratophore; inflated ocular areas lateral to base of
ceratophore with 2 pairs of eyes, the anterior pair larger than posterior
pair; upper tentacular cirri subequal in length to median antenna; lower
tentacular cirri shorter, subequal in length to ventral buccal cirri of seg-
ment II; lateral antennae short, subulate, on inner dorsal bases of ten-
tacular parapodia; capillary setae finely spinous and smooth; ciliated elon-
gate-conical tentacular lamellae medial to setal bundles (fig. la-d).
Neurosetae of segments II-[V with distal stems with variable number
spinous rows; blades elongate, slender, slightly hooked, with walls ir-

Genera Sthenelanella and Euleanira 433

0.5mm

fe)

0.1 mm

Fic. 3. Sthenelanella uniformis (Syntypes of S. atypica, USNM 32849):
a, Middle parapodium, anterior view; position of notopodial spinning
gland dotted in; b, middle parapodium, posterior view; c, neurosetae from
same; d, posterior parapodium, anterior view; e, posterior parapodium,
posterior view; f, neurosetae from same.

434 Proceedings of the Biological Society of Washington

regularly thickened on inner sides (figs. 1f, 2d); segment II] encroached
upon by segments II and IV, without dorsal cirri but with conical dorsal
tubercles and small branchiae; pair of small knobs or ctenidia dorsally,
medial to dorsal tubercles between sements II and III; additional pair of
ctenidia ventrally on segment III (fig. la, b, g).

Notopodia of parapodia forming conical acicular lobes and inflated
rounded upper lobes with ciliated ctenidia; two additional ctenidia be-
tween notopodia and branchiae (figs. 2,e,f; 3a-f). Notosetae capillary,
coarsely spinous, forming loose spreading bundles. Beginning on about
segment 14, notopodia provided with large oval spinning glands from
which slender threads emerge, extending far beyond the parapodia and
eventually becoming incorporated in their fibrous tough tubes. Neuropodia
diagonally truncate, with subequal rounded presetal and postsetal lobes;
presetal lobes with slight acicular notch. Neurosetae with blades short;
upper blades rod-shaped, rest conical, pointed. Ventral cirri short, sub-
ulate, with bulbous area on upper basal part and with terminal joint.
Pharynx with 2 pairs chitinous teeth and 13 or more pairs papillae (13 +
13; 14 + 15; 13 + 15). Tubes much longer than worms, branched, with
walls thick, tough, felted, covered with mud or sand (fig. 6, in Hartman,
1968).

Distribution: Southern California, Gulf of California to Ecuador. Lit-
toral to 73 meters, in silty, sandy, and muddy bottoms.

Remarks: The feltage notosetae and spinning glands were overlooked
by Moore (1910), but were observed by Hartman (1939) and the Berke-
leys (1941). Hartman (1961) questionably referred Sthenelanella atypica
to S. uniformis. The species is very common at shelf and slope depths off
southern California (Hartman, 1961).

Sthenelanella ehlersi (Horst), new combination

Figures 4, 5

Euleanira ehlersi Horst, 1916, p. 12, figs. 1, 2; 1917, p. 122, pl. 27, figs.
1-5.—Day, 1967, p. 101.

Material examined: Siboga station 2, Madura Strait, Dutch East Indies,
7° 25’ S, 113° 16’ E, 56 meters—2 Syntypes (ZMA 218); 1 Syntype
(RNHL 1190).

Description: As in generic diagnosis. Length up to 25 mm, width 2.5
min, including parapodia, and 3.5 mm, including setae; segments up to
75. Elytra delicate, transparent, on all segments from 25 on; first elytral
pair round, with few scattered sensory papillae on outer border; rest of
elytra subreniform to deeply sinuous on external border; anterior elytra
with transverse bands of brown pigment (fig. 4a,g,h). Prostomium with
ceratophore of median antenna equipped with lateral auricles on middle
of ceratophore; inflated ocular areas lateral to base of ceratophore with 2
pairs of eyes, the anterior pair larger than posterior pair; median antenna
and upper and lower tentacular cirri subequal in length; lateral antennae
short, digitiform, on inner dorsal bases of tentacular parapodia and addi-

Genera Sthenelanella and Euleanira 435

Fic. 4. Sthenelanella ehlersi (Syntype of Euleanira ehlersi, ZMA 218):
a, Anterior end, dorsal view, pharynx partially extended; first left and
first two right elytra removed; right palp missing; lateral antennae on
inner dorsal sides of segment I hidden from view; b, prostomium and ten-
tacular parapodium (I), lateral view; palp missing; c, parapodium of
segment II, posterior view; d, upper, middle, and lower neurosetae from
same (blade of middle ones missing); e, parapodium of segment III, an-
terior view; f, upper, middle and lower neurosetae from same; g, first two
right elytra; h, middle right elytron. (au, auricle; br, branchia; ct, ctenidia;
1 An, lateral antenna; t La, tentacular lamella).

436 Proceedings of the Biological Society of Washington

0.5mm
0.1 mm

Fic. 5. Sthenelanella ehlersi (Syntype of Euleanira ehlersi, ZMA 218):
a, Middle parapodium, anterior view; b, same, posterior view; c, upper,
middle and lower neurosetae from same; d, posterior parapodium, anterior
view; position of spinning gland dotted in; e, posterior parapodium, pos-
terior view; f, upper, middle and lower neurosetae from same.

Genera Sthenelanella and Euleanira 437

tional small oval ctenidia on their dorsal bases; elongate-conical tentacular
lamellae medial to setal bundles (fig. 4a, b). Neurosetae of segments II-
IV with distal stems with variable number spinous rows or these lacking;
blades elongate, slender, slightly hooked (fig. 4d, f); segment III without
dorsal cirri but with conical dorsal tubercles and small branchiae; pair of
small knobs or ctenidia ventrally (fig. 4e).

Notopodia of parapodia forming small rounded lobes with single prom-
inent ctenidia and slight indication of additional ctenidia (fig. 5a, b, d,
e). Notosetae capillary, finely spinous. Middle and posterior notopodia
provided with oval spinning glands from which long fine threads emerge
(fig. 5d, e). Neuropodia with rounded presetal and postsetal lobes, the
presetal lobes somewhat narrower. Neurosetae with short blades; upper
and lower blades rod-shaped; rest conical, pointed (fig. 5c, f). Ventral
cirri short, subulate, with bulbous area on upper basal part and with ter-
minal joint. Pharynx not extended and not examined. Tubes fibrous, stif-
fened with mud (Day, 1967).

Distribution: Dutch East Indies; South Africa (Natal). In 56 meters
{shallow to deep—Day, 1967).

Remarks: The spinning glands and long feltage notosetae were over-
looked by Horst (1916, 1917) but were observed by Day (1967), who
found specimens encased in fibrous tubes stiffened with mud. Due to the
opaque body, it is difficult to detect on which segment the spinning glands
begin. Day (1967) indicated that the elytra occurred on all segments from
21 on, instead of segment 25, as observed on the Syntypes.

KEY TO THE SPECIES OF STHENELANELLA

1. Anterior elytra with mottled pigmentation (fig. 2b); middle and
posterior elytra with external margins entire. Without oval ctenidia
on dorsal bases of tentacular parapodia (I) (fig. la). With pair
of oval ctenidia or knobs dorsally between segments II and III (fig.
|) ia no el aM canoe S. uniformis Moore

1’. Anterior elytra transversely banded (fig. 4a, g); middle and pos-
terior elytra with external margins deeply sinuous (fig. 4h). With
pair of oval ctenidia on dorsal bases of tentacular parapodia (1)
(fig. 4a, b). Without oval ctenidia or knobs dorsally between seg-
ments 11 and Ll {fig 4a) S. ehlersi (Horst )

Species of Sthenelanella are unique among the Sigalionidae in having
notopodial spinning glands which form notopodial threads that contribute
to their tough fibrous tubes, similar in this regard to some species of
Polyodontidae. Their neurosetae—all compound, with short blades—
sets them apart from most of the other species of Sigalionidae.

Sthenelanella polymorpha Hartmann-Schréder (1962), from Chile,
does not agree with the above diagnosis of Sthenelanella in a number of
characters, such as the relatively short palps, poorly developed tentacular
parapodia, and the shape of the parapodial lobes which lack spinous capil-
lary notosetae and spinning threads.

438 Proceedings of the Biological Society of Washington

LITERATURE CITED

BERKELEY, E. AND BERKELEY, C. 1941. On a collection of Polychaeta

Day, J. H.

from Southern California. Bull. So. Califormmia Acad. Sci., 40:
16-60, pl. 5.

1967. A monograph on the Polychaeta of Southern Africa.
Part 1. Errantia. Publ. Brit. Mus. (Nat. Hist.) London, No.
656: 1-458, 108 figs.

Hartman, 0. 1939. Polychaetous annelids. Pt. 1. Aphroditidae to Pisi-

onidae. Allan Hancock Pac. Exped. 1932-38, 7: 1-156, 28 pls.
1961. Polychaetous annelids from California. Allan Hancock
Pac. Exped., 25: 1-226, pls. 1-34.

1968. Atlas of the errantiate polychaetous annelids from Cali-
fornia. Allan Hancock Foundation Univ. So. California, Los
Angeles. Pp. 1-828, figs.

HARTMANN-SCHRODER, G. 1962. Zur Kenntnis des eulitorals der chilenis-

Horst, R.

chen Pazifikkiiste und der argentinischen Kiiste Siidpatagon-
iens unter besonderer Beriicksichtigung der Polychaeten und
Ostracoden,. Mitt. Hamburg. Zool. Mus. Inst., 60: 1-270, 223
figs.

1916. A contribution to our knowledge of the Sigalionidae.
Zool. Meded. Leyden, 2: 11-14, 2 figs.

1917. Polychaeta errantia of the Siboga-Expedition. Pt. 2.
Aphroditidae and Chrysopetalidae. Siboga-Exped. Leyden,
24b: 1-140, 5 figs., pls. 11-29.

Moore, J. P. 1910. The polychaetous annelids dredged by the U.S.S.

Albatross off the coast of Southern California in 1904; II.
Polynoidae, Aphroditidae and Segaleonidae. Proc. Acad. Nat.
Sci. Philadelphia, 62: 328-402, pls. 28-33.

Rersu, D. J. 1968. A biological survey of Bahia de Los Angeles, Gulf of

California, Mexico. II. Benthic polychaetous annelids. Trans.
San Diego Soc. Nat. Hist. 15: 67-106, 20 figs.

- 7
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ais 7
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4) afi re
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12 3 October 1969

PROCEEDINGS
OF THE

BIOLOGICAL SOCIETY OF WASHINGTON

PANDANUS DECUS-MONTIUM, A NEW SPECIES
FROM THE SOLOMON ISLANDS

By BENJAMIN C. STONE
School of Biological Sciences, University of Malaya,
Kuala Lumpur

Among the extensive collections of plants made in recent
years by the Forest Department of the British Solomon Islands
is a set of specimens representing the following new species of
Pandanus. Although as yet known only from staminate flower-
ing collections, the form of the leaves makes it very likely that
which includes Pandanus nemoralis Merr. & Perry P. paludosus
the species is a member of the Section Curvifolia B. C. Stone,
Merr. & Perry, and P. buinensis Merr. & Perry, all of which are
endemic to the Solomon Islands. They conform in having the
fruits (simple or 1-seeded drupes ) aggregated in cephalia, and
with horizontal rounded-reniform central stigmas, and vegeta-
tively in having rather broadly elliptic leaves, strongly nar-
rowed toward the base, which in life are downwardly curved
(hence the sectional name). The present newly proposed spe-
cies has, as may be seen from the accompanying illustrations,
leaves of this type, with a markedly narrowed and folded leaf-
base.

Although the species is here typified by a BSIP (British Sol-
omon Islands Plants) collection, I cite also specimens collected
by myself and L. J. Brass some years ago which were devoid of
flowers or fruit. Its striking appearance, at once noticeable in
the field, convinced me that it was new to science, but until
now no suitable (flowering) collections have been seen. It is
gratifying now to be able to describe this plant, which has been
named “decus-montium” (ornament of the mountains) to em-
phasize its attractive appearance.

33—Proc. Brot. Soc. WAsH., Vou. 82, 1969 (439 )

440 Proceedings of the Biological Society of Washington

Explanation of Figure 1

Fig. 1. Pandanus decus-montium new species.— Upper left: Habit
(Stone 2358, Malaita). Upper right: Habit (Stone 2358). Note markedly
flattened leaf-bases. Upper center: Staminal phalange, enlargement
(BSIP 1836). Lower left: large juvenile leaf, and small leaf from sucker-
shoot, of BSIP 1836 (LAE). Lower right: Staminate flowering branch,
with small leaves BSIP 1836 (LAE).

New Species of Pandanus 44]

Pandanus decus-montium new species (Sect. Curvifolia)
Fig. 1.

Arbor ad 10 m alta, stipite laeve cicatricata sparse ramosa, ramis erectis,
e basi pauci-radicante radicis gralliformibus ad 2 m altis. Folia anguste
oblanceolata usque ad 90 cm longa et 8 cm lata (in planta juvenili) vel
minora, 30 cm longa et 2.5 cm lata (in planta senili), apicem versus acuta
basem versus angustata et valde uniplicata pallidiora, marginibus apicem
versus crebre spinuloso-serratis, costis dorsaliter apicem versus similiter
spinuloso-serratis, basem versus inermibus vel subsparse aculeatis, aculeis
antrorsis 1 mm longis, plicis apicalibus ventraliter spinulosis. Lamina fol-
iorum supra viridis infra glauca basem versus pallida applanata. Inflore-
scentia foeminea ignota; mascula terminalis bracteata racemoso-spicata,
bracteis ca. 11, inferioribus foliaceis, basi navicularibus chartaceis, super-
ioribus toto navicularibus apice acuto costis marginibusque spinulosis.
Spicae masculae ovoideae ca. 3 cm longae et 1.5 cm latae albobrunneae.
Phalanges staminorum 5-8 mm longae, staminibus ca. 7-13 umbellatim
dispositis, filamentis fere 2 mm longis, antheris orbicularibus utrinque em-
arginatis 0.8-0.9 mm longis non apiculatis albidis.

Holotypus: BRITISH SOLOMON ISLANDS: GuapaLcANaL. Mount
Austen, alt. 1000 ft. “by stream on coral limestone; slender tree 30 ft tall
with conical crown; bole smooth, olive, with raised sinuous leaf-scars;
branches in twos or threes, upward-pointing, slender. Plants clumped.
Young stems unbranched and bearing much larger leaves in a very open
spiral; as stems branch their leaves get smaller. A few stilt roots to 6 ft.
Male tree, flowers fawn-color, subtended by brown papery bracts.” 9 May
1963, T. C. Whitmore BZIP. 1836 (BSIP); isotypes LAE! SING! K.

Additional specimens: BRITISH SOLOMON ISLANDS: Matarra.
Kwara-ae District, Kwalo, ridge about 1 mi. northeast of Tantalau Village,
ca. 1200 ft. alt. “Erect slender tree with terminal crown of large leaves and
a few lateral simple branches bearing small leaves; trunk somewhat tri-
quete-cylindric, ringed by subdistant leaf-scars and bearing scattered,
short, blunt prickles; base of trunk with several short, down-curved prop-
roots which are also prickly. Leaves in 3 distinct spirals, leaf-bases con-
spicuous by being flattened (folded); above the vase 6-8 inches each
leaf margin is then flattened horizontally; the M-shape of most pandan
leaves is scarcely noticeable. No flowers or fruit seen.” 23 September
1957, B.C. Stone 2358 (BISH!).

SAN CRISTOVAL. Hinuahaoro, 900 m. alt., “tree to 10 m. high, stems
solitary, or several erect from a curved or horizontal common stock, raised
several feet from the ground on stout prickly proproots. Stem with numer-
ous lateral small upturned branches. Leaves not over 1 m. long, glaucous
beneath, of soft texture, numerous in 3 spirals. Sterile.” 22 September
1932, L. J. Brass 2918 (A!).

Vernacular names: “apapola” (BSIP 1836); “ahole” (Stone 2358).

I would like to acknowledge the helpful advice of Dr. T. C. Whitmore
and of G. F. C. Dennis in connection with the preparation of this paper.

442. Proceedings of the Biological Society of Washington

2 3 October 1969

PROCEEDINGS
OF THE

BIOLOGICAL SOCIETY OF WASHINGTON

USE OF MALE EXTERNAL GENITALIC DETAILS AS
TAXONOMIC CHARACTERS IN SOME SPECIES OF
PALAEMONETES (DECAPODA, PALAEMONIDAE )!

By LaurENCE E. FLEMING
Mississippi State University?

The various species of shrimp belonging to the genus Palae-
monetes are currently identified and classified on the basis of
the spinose ornamentation of several different body sites.
Because of overlap and variation, identification and relation-
ships are difficult to determine.

The taxonomic characters currently in use for Palaemonetes,
with the possible exception of cheliped proportions, are prob-
ably subject to ecological modifications. This is especially true
of spinose ornamentation. On the other hand, the structures
utilized in amplexus in most Crustacea are usually stable and in
so far as is known, are little affected by ecological factors. This
is especially true in many Malacostraca where male appendages
have been particularly useful in taxonomic and evolutionary
studies.

In this study the external genital details of the second
pleopod of male Palaemonetes were examined and evaluated as
taxonomic characters for identification. The study was limited
to the epigean species of the Gulf Coastal Plain of the United
States and its coastal waters. Stable, and unique for the spe-
cies, setal ornamentation of the tip of the appendix masculina
of the second pleopod and equally stable gross morphology of
the second pleopod provide potentially valuable characteristics
for the systematic study of Palaemonidae.

1 This research was supported in part by NSF Grant No. GB-4719 to J. F.
Fitzpatrick, Jr.

2 Present address: Department of Biology, Virginia Polytechnic Institute, Blacks-
burg, Virginia.

34—Proc. Biot. Soc. WAsH., Vou. 82, 1969 (443)

444 Proceedings of the Biological Society of Washington

Species Studied: Holthuis (1949, 1952) listed six epigean species of
Palaemonetes as occurring in the United States, and five of these are
known from the state of Mississippi and adjacent waters (ibid.): P. (P.)
kadiakensis Rathbun, 1902; P. (P.) paludosus (Gibbes, 1850); P. (P.)
vulgaris (Say, 1818); P. (P.) pugio Holthuis, 1949; P. (P.) intermedius
Holthuis, 1949. Specimens of these species from Mississippi were the
principal source of information although most specimens of P. paludosus
were collected in St. Charles and St. James parishes, Louisiana.

Additional material was available from various sources, and specimens
from four Gulf states (Mississippi, Louisiana, Texas, and Alabama) were
analyzed. Material studied included five collections from two counties in
Alabama, 21 collections from five parishes in Louisiana, two collections
from two counties in Texas and 42 collections from 15 counties in Missis-
sippi.

Following initial evaluations, type specimens of these species at the
Smithsonian Institution were examined and the conclusions validated.
Other types at SI were also examined for the same characteristics to deter-
mine if the proposed criteria could be applied to other species of the ge-
nus. These species, studied by means of type material at SI, are: P. (P.)
schmitti Holthuis; P. (P.) suttkusi Smalley; P. ( Alaocaris) antrorum Bene-
dict. Type material of P. (P.) eigenmanni Hay and P. (P.) hiltoni
Schmitt were also examined at SI, but the pleopods were disarticulated,
shriveled or in some way damaged and little use could therefore be made
of this material.

Procedures: Specimens of Palaemonetes were collected by means of dip
nets, seines, and on occasion, use was made of boat-drawn trawls and
plankton nets. The specimens were collected from large and small ponds,
roadside sloughs, running streams, inlet bayous of the Mississippi Sound
of the Gulf of Mexico and from the Gulf of Mexico itself. Freshwater
species were found to be the most abundant in floating, submerged and
emergent aquatic vegetation such as duckweed (Spirodela sp. and Lemna
sp.), water hyacinth ( Eichhornia sp.), pondweed ( Potamogeton spp.) and
bladderwort (Utricularia sp.). Brackish and marine species were mostly
obtained from growths of eelgrass ( Vallisneria spp. ).

A total of 1582 specimens were studied in detail: 845 specimens of P.
kadiakensis, 169 specimens of P. paludosus, 430 specimens of P. pugio, 71
specimens of P. vulgaris, and 67 specimens of P. intermedius.

Individuals were randomly selected and the second left pleopods re-
moved and temporarily mounted on slides using Monk’s Mounting Me-
dium (5cc corn syrup, 5cc fruit pectin, 3cc of water with thymol added as
a preservative). Each pleopod was placed in precisely the same position
on the slide. Then a careful examination of each pleopod was made using
a compound microscope. To further validate the conclusions drawn, sev-
eral pleopods were dismounted from their position on the slide, rearranged
in exactly the opposite position, and again subjected to microscopic ex-
amination.

Concluding these studies, camera lucida drawings were made of selected

Decapod male genitalia 445

representative pleopods. Two sets of drawings were made of each. One
drawing was made under the low power (35x ) of a compound microscope
to reveal the structure of the entire pleopod, and another drawing was
made under the high power (450 ) of the tip of the appendix masculina
of each male specimen. The coverslips of the slides were then ringed with
“Permount’ to insure their permanency.

Following this, in situ camera lucida drawings were made of the second
right pleopod of randomly selected specimens using a magnification of
38x on a stereoscopic microscope. This was done to validate the accuracy
of the orientations and interpretations of disarticulated appendages.

OBSERVATIONS

In all of the specimens examined, the general appearance of the ex-
ternal secondary sex characters of the five different species seem to re-
semble Meehean’s (1936) description of those of P. paludosus (probably
P. kadiakensis). In Palaemonetes the first two pairs of pleopods exhibit
sexual dimorphism and modification. The first pair of pleopods (which
are not copulatory organs) are relatively unmodified but the inner ramus
(= endopodite proper) of the male is approximately three times as long as
that of the female. In both sexes the endopodite and exopodite are heavily
armed with plumose setae. The second pair of pleopods (the pair exhibit-
ing the greater modification) have the endopodite modified to include
a non-podomere appendix interna (= retinaculum of Meehean, 1936)
arising from the inner margin in both sexes (Figs. 1 and 2). Between the
endopodite proper and the appendix interna of the male an accessory
process arises near the base of the appendix interna and is termed the
appendix masculina (op. cit.).

Interspecific variability of the second pleopod of male specimens of
Palaemonetes were found. The gross morphology of the various parts of
the pleopod varies in such factors as proportional lengths of certain parts,
shape of the tip of the appendix interna and overall shape of the appendix
masculina. Another species specific feature is the setal ornamenation of
the tip of the appendix masculina. The number of apical setae and the
number of subapical setae are sufficient to identify most species, except P.
vulgaris and P. intermedius which are identical in this respect.

P. intermedius (Fig. 3) is used as an example to illustrate the position
of the apical setae in reference to the subapical setae. There are four apical
setae and two subapical setae in P. intermedius and P. vulgaris (Fig. 4).
There are three apical setae and one subapical seta in P. kadiakensis (Fig.
5); four apical setae and one subapical seta in P. paludosus (Fig. 6); five
apical and one or two subapical setae in P. pugio (Fig. 7).

In summary, the setal ornamentation of the appendix masculina is
specifically unique in the species of Palaemonetes examined, with the one
exception, as stated above, involving P. vulgaris and P. intermedius. All
the other species are easily separable using this character alone.

Study of the gross morphology of the pleopod reveals differences in pro-
portions of two structures: (1) extent of the appendix interna along the

446 Proceedings of the Biological Society of Washington

5

APICAL SETAE————(----=
SUBAPICAL SETAE ;

APPENDIX MASCULINA

Fics. 1-6. Second pleopod of various species of Palaemonetes. 1. Mesial
view of P. kadiakensis Rathbun illustrating gross morphology of a female
pleopod. 2. Mesial view of P. paludosus (Gibbes) illustrating gross mor-
phology of a male pleopod. 3. Tip of appendix masculina of P. inter-
medius Holthuis showing setal ornamenation. 4. Tip of appendix mas-
culina of P. vulgaris (Say) showing setal ornamentation. 5. Tip of ap-
pendix masculina of P. kadiakensis Rathbun showing setal ornamentation.
6. Tip of appendix masculina of P. paludosus (Gibbes) showing setal or-
namentation.

Decapod male genitalia 447

12

Fics. 7-14. Second male pleopod of various species of Palaemonetes. 7.
Tip of appendix masculina of P. pugio Holthuis showing setal ornamenta-
tion. 8. Mesial view of P. kadiakensis Rathbun illustrating specific gross
morphology; exopodite concealed. 9. Mesial view of P. pugio Holthuis il-
lustrating specific gross morphology. 10. Mesial view of P. vulgaris (Say)
illustrating specific gross morphology. 11. Mesial view of P. intermedius
Holthuis illustrating specific gross morphology. 12. Mesial view of P.
schmitti Holthuis (from the Canal Zone) illustrating specific gross mor-
phology. 13. Mesial view of P. suttkusi Smalley (from Mexico) illustrat-
ing specific gross morphology. 14. Mesial view of P. antrorum Benedict
(a subterranean species) illustrating specific gross morphology.

448 Proceedings of the Biological Society of Washington

appendix masculina and (2) extent of the appendix masculina along the
endopodite proper.

The appendix interna extends along slightly more than the proximal one-
half of the appendix masculina in P. kadiakensis (Fig. 8); along less than
the proximal one-half of the appendix masculina in P. paludosus (Fig. 2);
along the proximal three-fourths of the appendix masculina, and in some
specimens the appendices may even be subequal in P. vulgaris (Fig. 10).
In P. pugio (Fig. 9) and P. intermedius (Fig. 11) the appendix interna
extends along the proximal two-thirds of the appendix masculina. Through
the use of this character all species except P. pugio and P. intermedius can
be distinquished.

The appendix masculina extends to the distal one-third of the endopo-
dite proper in P. vulgaris (Fig. 10) and P. pugio (Fig. 9); to the distal
one-fourth of the endopodite proper in P. intermedius (Fig. 11); to the
distal one-eighth of the endopodite proper in P. kadiakensis (Fig. 8); to
the distal one-tenth of the endopodite proper, and in some specimens the
two appendices are subequal in P. paludosus (Fig. 2). Thus, P. vul-
garis and P. pugio are alike in this character.

The shape of the tip of the appendix interna in P. paludosus (Fig. 2) is
large and flat or paddle-shaped while in all other species it is small and
round.

The overall shape of the appendix masculina differs in two species from
the shape of the structure in the other three species. In P. kadiakensis
(Fig. 8) and in P. pugio (Fig. 9) the appendix masculina is gently curved
laterad, with the curve in both species occurring in the approximate
area of the limit of extent of the appendix interna along the appendix
masculina. In P. paludosus (Fig. 2), P. vulgaris (Fig. 10), and P. inter-
medius (Fig. 11) the appendix masculina is straight. In P. paludosus the
appendix masculina is straight and stiffened in comparison with the ap-
pendix masculina of P. vulgaris and P. intermedius.

A summary of the various features of the second pleopod of these five
species is given in Table 1. The types of all are comparable with the fig-
ures (Figs. 1-11) and Table 1.

Type material from areas outside of the Gulf Coastal Plain was ex-
amined at SI and found to exhibit species unique characteristics. In P.
schmitti (Fig. 12.) there is a very complicated setal ornamentation (eight
long apical setae); one short subapical seta; and the appendix interna turns
upward and outward. In P. suttkusi (Fig. 13) there are five very short
apical setae arranged in a circle; one short subapical seta; and the ap-
pendix interna is very short and straight. In P. antrorum, a subterranean
species, (Fig. 14) there are two long apical setae; three long subapical
setae; the appendix interna is relatively straight; and the appendix mas-
culina is very stout.

DiscussIONn

Although Holthuis’ revision (1952) of the subfamily Palaemoninae now
serves as the primary source for all taxonomic work in this group, specific,

Decapod male genitalia 449

TABLE 1. Characteristics of second pleopod of certain species of

Palaemonetes.
Appendix masculina Appendix interna
Num-
Num- ber of Extent along
ber of sub- Shape endopodite Shape Extent along
apical apical of proper of appendix
Species setae setae tip tip masculina
To slightly more
kadiakensis 3 1 curved proximal small, round than
1k proximal 14
To
paludosus 4 1 straight proximal large, flat or less than
%o paddle-shaped proximal 14
To
pugio 5 1-2 curved proximal small, round equal to
% proximal 2%
To slightly more
vulgaris 4 2 straight proximal small, round than
x proximal 34
To
intermedius 4 22 straight proximal small, round equal to
34 proximal %
To slightly more
schmitti 8 1 straight proximal small, round than
% proximal 34
To
suttkusi 5 1 curved proximal small, round less than
% proximal 14
To
antrorum 2 3 straight proximal small, round equal to
% proximal %

subgeneric and generic characters utilized by him exhibit overlap, variabil-
ity and, in some cases, indistinctiveness which tend to make the identifica-
tion of certain specimens extremely difficult. Some characters are difficult
to evaluate. For example, the pleura of the fifth abdominal segments are
used as a diagnostic character in the separation of P. pugio, P. vulgaris,
and P. intermedius yet these features are intraspecifically mutable.

The use of the second pleopod of the male as a source of taxonomic
characters in Palaemonetes would alleviate many difficulties now en-
countered in identifying species of Palaemonetes and in so doing would
make a contribution to the taxonomy of the group. Further, evolutionary
and interspecific relationships probably can be better evaluated with more
stable characters.

Holthuis (1952) noted that P. vulgaris, P. pugio and P. intermedius are
very similar and for a long time have been confused under the name P.
carolinus and P. vulgaris. He listed the characters that he used to separate

450 Proceedings of the Biological Society of Washington

them and then concluded by stating that, although the three species can be
generally easily separated, there are examples, especially with juveniles,
when identification of a specimen is difficult. I should note that the use of
details of the male external genitalia in identification does not remove the
difficulties in identifying juveniles and females; its value lies in the in-
creased confidence its use imparts to identifications of adult male speci-
mens.

The separation of P. vulgaris from P. intermedius is extremely difficult
with the use of the characteristics of the appendix masculina. Differentia-
tion between the two, in fact, can be attained solely through the use of
the proportional lengths of the appendix interna and appendix masculina.
Further, the significance of these two characters has yet to be determined,
especially with reference to other characteristics of the two populations.

There is similarity in larval development of P. pugio, P. intermedius,
and P. vulgaris (Broad, 1957; Broad and Hubschman, 1962). Broad and
Hubschman (1963) working with P. kadiakensis larvae and Dobkins
(1963) working with P. paludosus larvae found that the larval develop-
ment of the freshwater forms could be distinquished from the salt
water forms by the tendency toward condensation of the stages, larger
eggs and larger larvae of the freshwater forms.

From this information, together with that gathered from studies on
the male second pleopod, one concludes that P. pugio, P. vulgaris, and P.
intermedius are indeed related. P. pugio, however, is conspicuously
different and easily separable, and thus specifically distinct from either P.
vulgaris or P. intermedius. P. vulgaris and P. intermedius, on the other
hand, are visibly identical in such a varying array of features and percep-
tibly different in so few characters that they could be conspecific. This
view cannot be supported satisfactorily or refuted until more information
is provided, particularly detailed populational and ecological data. For
the time being, at least, there are distinct morphological differences be-
tween the two and there are no evidences of interbreeding. Therefore,
they are best considered specifically distinct. P. kadiakensis and P. pal-
udosus are clearly morphologically distinct from each other and from
the other species. Thus, they, too, represent separate species. Examination
of type material at the United States National Museum confirmed the dis-
tinctiveness of the morphological features of the gential apparatus of the
male second pleopod of the five species of Palaemonetes studied.

Epigean species of Palaemonetes from areas outside the Gulf Coastal
Plain of the United States apparently also are specifically unique in these
characteristics as revealed by type specimens from Mexico and the Canal
Zone examined at SI. Subterranean species of Palaemonetes probably also
may be similarly identified if studies made on type material of Palae-
monetes ( Alaocaris) antrorum Benedict, 1896, at SI are any indication.

SUMMARY

1. The second pleopod of the male may be used in taxonomic evaluation
of epigean species of Palaemonetes.

Decapod male genitalia 451

2. P. kadiakensis, P. paludosus and P. pugio can be easily separated from
one another, and all can be differentiated from P. vulgaris and P.
intermedius using counts of apical and subapical setae of the tip of the
appendix masculina; P. vulgaris and P. intermedius cannot be distin-
quished using this character.

3. The relative length of the appendix interna can be used to distinquish
P. kadiakensis, P. paludosus and P. vulgaris, although P. pugio and P.
intermedius are indistinguishable by this character.

4. The relative length of the appendix masculina is distinctive in P. pal-
udosus, P. kadiakensis and P. intermedius, but similar in P. vulgaris
and P. pugio.

5. The tip of the appendix interna is large and flat or paddle-shaped in
P. paludosus; it is small and round in all other species.

6. The appendix masculina is curved laterad in P. kadiakensis and P.
pugio and straight in all other species; it is decidedly stiffened in P.
paludosus.

7. Studies of the second pleopod and statements of other students sug-
gest that P. intermedius and P. vulgaris may be conspecific, but they
are considered here as distinct species; all other species are clearly dis-
tinct.

8. Studies made on type specimens of P. schmitti, P. suttkusi and P.
antrorum at USNM revealed that they likewise may be separated from
each other and from other species by these criteria.

ACKNOWLEDGMENTS

The writer wishes to express his sincere appreciation to Dr. J. F. Fitz-
patrick, Jr., for his assistance and supervision of this study and for review-
ing the manuscript. Dr. Fitzpatrick was not only an invaluable counsel
throughout the course of this research but a constant source of encourage-
ment. Dr. Perry C. Holt, Virginia Polytechnic Institute, carefully read the
manuscript and made helpful suggestions. Drs. Gordon Gunter and Walter
Abbott generously allowed use of facilities at the Gulf Coast Research
Laboratory, and Mr. J. Y. Christmas and Mr. G. P. Garwood made mu-
seum specimens available; Mrs. Shirley Dimmick provided numerous aids
in securing literature and information on Palaemonetes. Dr. James Frank-
lin Payne and Mr. Shih-ming Chien were generous with help in collect-
ing specimens. Dr. M. Saeed Mulkana is especially thanked for his help
and assistances far too numerous to enumerate. Dr. Roger F. Cressey, Di-
vision of Crustacea, Museum of Natural History, Smithsonian Institution,
kindly made it possible for me to study type specimens of P. schmitti, P.
suttkusi and P. antrorum.

LITERATURE CITED

Broap, A. C. 1957. Larval development of Palaemonetes pugio Holthuis.
Biol. Bull. 112: 144-161.
AND J. H. HuspscumMan. 1962. A comparison of larvae and
larval development of species of eastern United States Palae-

452 Proceedings of the Biological Society of Washington

monetes with special reference to the development of Palae-
monetes intermedius Holthuis. Amer. Zool. 2: 394-395 ( Ab-
stract).
1963. The larval development of Palaemonetes kadiakensis
M. J. Rathbun in the laboratory. Amer. Microscop. Soc., Trans.
82: 185-197.

Doskin, S. 1963. The larval development of Palaemonetes paludosus
(Gibbes, 1850) (Decapoda, Palaemonidae), reared in the lab-
oratory. Crustaceana, 6: 41-61.

Grspes, L. R. 1850. On the carcinological collections of the United
States. Proc. Amer. Ass. Advance. Sci. 3: 167-201.

Ho.tuuts, L. B. 1949. Note on the species of Palaemonetes (Crustacea,
Decapoda) found in the United States of America. Proc.
Konink. Nederland Akad. Wetensch. 52: 87-95.

. 1952. A general review of the Palaemonidae (Crustacea,

Decapoda, Natantia) of the Americas. II. The subfamily Pal-
aemoninae. Occ. Pap. Allan Hancock Found. 12: 1-396.

MEEHEAN, O. L. 1936. Notes on the freshwater shrimp Palaemonetes
paludosa (Gibbes). Amer. Microscop. Soc., Trans. 55: 433-
44],

RaATHBUN, M. J. 1902. Descriptions of new decapod crustaceans from the
West Coast of North America. Proc. U.S. Nat. Mus. 24: 885-
905.

Say, T. 1818. An account of the Crustacea of the United States. J.
Acad. Nat. Sci., Philadelphia 2: 235-458.

Vol. | 3 October 1969

PROCEEDINGS oy
OF THE f ne 790 19869
BIOLOGICAL SOCIETY OF WASHINGTON. - |

CONTRIBUTIONS TO A REVISION OF THE
EARTHWORM FAMILY LUMBRICIDAE
V. EISENIA ZEBRA MICHAELSEN, 1902!

By G. E. Garters
Zoology Department, University of Maine, Orono

Twenty species of lumbricid earthworms, brought by man
from Europe since 1500 A. D. (Gates, 1966 and 1967), have
been found to be variously domiciled in North America. Another
species, that may eventually become widely distributed through-
out the continent, is now added to the list. At least one more,
perhaps others, will be added later.

American material and data were supplied by Mr. Salvatore
Billeci. For comparison, an identified series of the same species
from Wales was provided by Dr. K. Sylvia Richards.

Eisenia Malm, 1877 (emend. Gates, 1968 )
Eisenia zebra Michaelsen, 1902

Eisenia veneta var. zebra Michaelsen, 1902. Mitt. Naturhist. Mus. Ham-
burg, 19, p. 39 (Type locality, Chosta, Kreis Sotschi, Transcaucasia.
Type, supposedly in the St. Petersburg Mus. )

Helodrilus (Eisenia) venetus var. zebra, Michaelsen, 1910. Ann. Mus.
Zool. Acad. Sci. St. Petersburg, 15, p. 3.

Dendrobaena veneta var. zebra, Pop, 1943. Ann. Hist. Nat. Mus. Hun-
garici, (Zool.), 36, p. 22. Brinkhurst, 1962, Proc. Zool. Soc. London,
138, p. 325. Gerard, 1964, Linnean Soc. London, Synopses British
Fauna, No. 6, p. 39, etc.

Material examined: San Francisco, California, 1-3-20(-+), received on
several occasions from S. Billeci. Identified specimens, from Britain, 1-1-
32, provided by K. Sylvia Richards.

External characteristics: Size, 51-96 by 5 (an aclitellate) to 8 mm. Seg-
ments, 83-153 (Table), 113-153 (unamputated specimens ). The majority
of the unamputated worms have segment numbers in the range of 127-
147. The average for 44 unamputated specimens (Table, Nos. 10-59 but

1 From research financed by the National Science Foundation.

35—Proc. Biot. Soc. WasuH., Vou. 82, 1969 (453 )

ee”

454 Proceedings of the Biological Society of Washington

Typhlosole termination and segment number in Eisenia zebra

Typhlosole Atyphlo- Soma Typhlosole Atyphlo- Soma
Serial ends in solate seg- Serial ends in solate seg-
number segment segments ments number segments segments ments
1 78 5 83 31 121 16 137
2 79 4 83 32 121 19 140
3 83 4 87 33 122 14 136
4 88 vi 95 34 122 15 137
5 93 10 103 35 122, AGT 139
6 94 6 100 36 123 13 136
7 94 8 102 37 123 14 137
8 98 8 106 38 123 16 139
9 98 1] 109 39 123 17 140
10 101 12 113 40 125 15 140
11 106 10 116 41 125 16 141
12 109 8 117 42 126 8 111
13 109 11 120 43 126 16 142,
14 109 12 121 44 128 14 142
15 110 16 126 45 128 15 143
16 110 18 128 46 128 15 143
17 111 15 126 47 129 15 144
18 112 16 128 48 130 4 134
19 112 18 130 49 130 8 138
20 114 15 129 50 130 14 144
21 117 15 132 51 130 15 145
22 117 19 136 52 130 16 146
23 118 16 134 53 130 16 146
24 118 17 135 54 131 16 147
25 119 8 127 55 131 17 148
26 119 15 134 56 132 16 148
27 119 18 137 57 135 8 143
28 120 8 128 58 137 10 147
29 120 15 135 59 137 16 153.
30 120 18 138
NOTES

Worms numbers 2, 9, 14, 17 were posterior amputees having each an obviously
regenerated periproct. ,

The typhlosole of No. 12 was rudimentary in the 108th—109th segments, having
been reduced after posterior amputation.

Those of Nos. 1-9 not already mentioned probably were old posterior amputees.
Those numbered 25, 28, 42, 49, 57 are believed to be posterior amputees.

Coelomic cavities of the last few segments in Nos. 40 and 41 were filled with
brown bodies of various sizes and shapes.

excluding 12, 28, 42, 48, 49, 57) is 136.5. The mean number of segments
for 47 specimens is 136.4681, with a standard deviation of 8.5539 and a
standard deviation of the mean of 1.2477. A majority of the worms have
segments in a range of 134-148, which is about the size of the range for
the majority of E. hortensis (Gates, 1968b). Color, dark red to slate, in

Earthworm revision 455

transverse bands, leaving a fairly wide uncolored band centered at each
intersegmental furrow, sparse in some portion of the dorsum in ix—x. Soma,
posteriorly almost transversely rectangular in cross section with b and d
setae at the four corners. Prostomium epilobous, tongue open (all). When
the pre-oral lobe is drawn more or less completely inside the buccal cavity,
the lobe is demarcated from the tongue by a transverse furrow. Such a
condition is called “combined pro- and epilobous”. Several worms were
almost tanylobous. The periproct often is large, with an anterior portion
showing evidence of differentiation of another metamere, such as presence
of a dorsal pore (which may not be functional) or presence of setae more
anteriorly. If both conditions are recognizable the area was counted as two
segments even though not yet demarcated from each other by an inter-
segmental furrow. Secondary annulation, lacking.

Setae, present from ii, widely paired, CD ca. = or very slightly < AB,
BC slightly < or ca. = AA < DD < %4C. Nephropores, inconspicuous,
actual pores never seen, locations occasionally recognized, probably at B in
xiv, xv usually, at or above D in first few segments, elsewhere varying ir-
regularly and with asymmetry between a ventral level just above B and a
dorsal level above D. First dorsal pore, at 4/5 (1 specimen), 5/6 (35),
pores at 9/10, 10/11 of clitellate individuals occluded.

Clitellum, saddle-shaped, reaching down below C, dorsal pores oc-
cluded, intersegmental furrows not obliterated, (xxvi)—xxxiii (4), xxvi-
xxxiii (29'), xxvi/eq-xxxiii (1), xxvii—xxxiii (7), (xxvii)—xxxiii (1), xxviii-
xxxiv (1). Tubercula pubertatis, longitudinally placed, often bounded
laterally by a distinct furrow, median borders often uncertain and seem-
ingly just including b setae, anterior and posterior borders usually uncer-
tain, xxix—xxxi (10), xxix/eq-xxxii/eq (10), xxix/eq-xxxii (4), xxx—xxxi
(14), xxx—xxxii/eq (1), xxx—xxxii (4). Sometimes a nearly circular por-
tion in each of xxx and xxxi seems more prominent.

Genital tumescences, slight, boundaries very indistinct, including some
or all of the setae in ix or xii, a,b separately in xxviii-xxxi (2), xxviii-xxxii
(3), xxix—xxx (3), xxix—xxxi (3), xxix—xxxii (17).

Internal anatomy: Septa, 5/6—-12/13 slightly strengthened, 13/14-
15/16 muscular and increasingly thickened posteriorly.

Special longitudinal muscle band at mD, present from 5/6. Pigment,
red, in circular muscle layer. Peritoneum blistered away from musculature
in dorsum of ix—xi. Broad transverse stripes of pigment sometimes are as-
sociated with the dorsal peritoneum anteriorly.

Calciferous sacs and lamellae, lacking in x. Esophagus widest in xi—xii
where the lumen is narrow. Usually no marked external constriction at in-
sertion of 11/12. Lamellae are largest in xi in which segment the gut
always is whiter or redder than anteriorly or posteriorly. Esophageal
valve, in xiv (35). Intestinal origin, in xv (35). That portion of the in-
testine belonging in xv occasionally has been drawn back into gut lumen
of xvi. Gizzard, mostly in xvii, but fenestration dorsally of 17/18 and
18/19 contributes to an appearance of greater posterior extent. Typhlo-
sole, present from region of xxii, at first with widened and flat ventral

456 Proceedings of the Biological Society of Washington

face. A cross section at first has an inverted T-shape but subsequently is
obviously though only slightly bifid. The typhlosole ends as shown in the
table but, when there was no posterior amputation, in region of the 101st
to 137th segments, usually in the 110th—130th. Up to 18 intestinal seg-
ments were atyphlosolate.

Dorsal blood vessel, single, recognizable forward only to 5/6. How-
ever, in one worm, a small section of the trunk among the pharyngeal
glands was blood-filled and traceable to a bifurcation under the brain.
Ventral trunk, complete, bifurcating over subpharyngeal ganglion. Sub-
neural trunk, complete, bifurcating at anterior end of the nerve cord, ad-
herent to cord but when distended coming easily away. Extra-esophageal
trunks, median to hearts, turning up to dorsal trunk in xii (35). Hearts,
in vii—xi (35:), none seen in vi.

Nephridia, vesiculate. Bladders, elongately sausage-shaped, transversely
placed in BD or reaching beyond D, joined at lateral end by looped tu-
bule, narrowing as they pass downward and into parietes close to B but
without a distinct duct.

Holandric. Testes and male funnels free in coelomic cavities. Male
funnels, polyplicate, sometimes complexly so and then rosette-like. Male
gonoducts, without epididymis (35), passing straight laterally to parietes,
disappearing from sight in an anterior portion of the atrial glands in xiv.
Seminal vesicles, 4 pairs, smallest in x, the last pair largest and at height of
maturity extending in posterior pockets of 12/13 back to level of 14/15.

Ovaries, each with a terminal egg string that may contain 4-7 ova (35).
Ovisacs, present in xiv (35). Spermathecae, in ix and x (35), each with
a short and slender but definitely coelomic duct. Ampulla, spheroidal to
ovoidal, occasionally more or less reniform to almost bilobed.

TP glands, acinous, more or less conspicuously protuberant into the
coelomic cavities, just lateral to B. Atrial glands, acinous, usually entirely
within the body wall which is markedly bulged into the coelomic cavities
of xiv—xvi in the median portion of BC. An equatorial cleft is obvious in
xv. Setal follicles of ix (2), xii (24) are enlarged, conspicuously protuber-
ant in the coelom, each surrounded by a rosette of acinous supraparietal
glands. Setae of those follicles are of the usual genital sort. The body
wall ventrally in BB of xvi-xxiii is blistered away from the musculature
and the space between peritoneum and muscles is filled with a delicate
coagulum. GS glands were not certainly distinguished among the blisters.

Reproduction: Spermatozoal iridescence on male funnels of clitellate
worms showed that maturation of sperm had been completed. Iridescence
in the spermathecal ampullae proved the worm had copulated. In absence
of any contra-indication, reproduction accordingly can be assumed to be
amphimictic.

Some of the spermatophores obtained were found to contain sperm.

Distribution: Outside of Russia, E. zebra had been found in Turkey,
Wales, England, Ireland, but records for extra-Russian areas are few.

Cocoons: Color, a light lemon-yellow, perhaps becoming brown later.
Shape, tapering slightly at each pole to a protuberance. One terminal

Earthworm revision 457

protuberance usually is markedly thicker than the other but length and
shape of each free end vary considerably. Micrometer measurements sup-
plied by Mr. Billeci are as follows: Diameter, at thickest equatorial
portion, 0.1415, 0.143, 0.144, 0.145 (twice), 0.146 inches. Average of the
six measurements, 0.144 inch. Length, exclusive of the polar appendages,
0.125, 0.126, 0.138, 0.154, 0.155 (twice), 0.158 (twice), 0.159,
0.165, 0.186, 0.201 inches. Average of twelve cocoons, 0.1568 inch.

Spermatophores: One was noted on each of four worms, two were seen
on each of two worms. Always discoidal and transparent, shape varied
from subcircular to elliptical. Each had a small, opaque central thicken-
ing that contained sperm. Locations: across 27/28, in AB or centering at
B, or extending across all of xxviii-xxix and centering at A.

Autotomy: No. 1 had half completed breaks in body wall, on left side
only, at 102/103 and 93/94. No. 2 had a half completed break on left
side at 80/81. No. 3 had completed a break on the right side at 89/90. No.
4 had a completed break in ventral body wall only at 88/89, but at 103/
104 the break had been completed—the parts held together only by
cuticle.

Regeneration: Absence of head and tail regenerates, except for four
periprocts, in a total of more than 60 specimens seems unusual especially
in comparison with its relative Eisenia foetida (Savigny, 1826).

Abnormality: Three spermathecae were present in x (1).

Parasites: Long nematodes were present in the ventral blood vessel (2
hosts ). Seminal vesicles of ix,x (1 worm) were filled with small cysts and
similar cysts were present in coelomic cavities of x—xi.

Remarks: From an anterior portion dorsally of a worm that was about
to be put on hook there came out a creamy yellow fluid. The liquid,
which may have been from distended spermathecae, had a strong odor, ac-
cording to Mr. Billeci and two of his party, like that of decaying bananas.

SYSTEMATICS

Eisenia veneta (Rosa, 1886) has been at one time or another in five
lumbricid genera, Allolobophora, Bimastos, Dendrobaena, Eisenia, Helo-
drilus. At present some European specialists place the species in Dendroba-
ena while others refer it to Eisenia.

Eisenia, recently redefined (Gates, 1968a) in accordance with conserva-
tive somatic anatomy, lacks calciferous sacs. The calciferous gland opens
directly, i.e., without intervention of sacs, into the gut lumen in xi. The
calciferous gland of specimens identified by Michaelsen as the typical form
of “Helodrilus venetus” was studied by Smith (1924, p. 27). He stated
that the anterior end of the gland was in x as Omodeo (1952, p. 190) also
thought. Actually a portion of the gut belonging in xi had been herniated
into x as Omodeo later (1954, p. 128) discovered.

Insofar as the calciferous gland and “var. typica” are concerned Rosa’s
veneta probably can go in Eisenia. However, confirmation is required
from other somatic anatomy.

Sixteen varieties of E. veneta were given Latin names by European

458 Proceedings of the Biological Society of Washington

zoologists. Some still are in use. One, var. hortensis Michaelsen, 1890,
now more adequately characterized (Gates, 1968b) is recognized as a
species. Another, var. hibernica Friend, 1893, subsequently will be shown
to be distinct. Other taxa are distinguishable at present from each other,
if at all, only by characters of dubious systematic value (cf. Omodeo,
1952, p. 8 and/or Gerard, 1964, p. 38-39). The definition of “f. typica”
(Gerard, 1964, p. 38) almost covers the entire range of variation in all
varieties. By 1893 Rosa himself already had referred to his “veneta” in-
dividuals with a clitellum extending from xxiv, xxv, xxvi, or xxvii through
XXxiii Or Xxxiv.

Information as to existence of types of Rosa’s veneta, as well as material
of various varieties (including f. or var. “typica’”), has been unobtainable.
Michaelsen’s variety, so far as can be discovered from the literature, has
been recognized on several occasions without difficulty. Populations
from which the present samples were obtained seemingly are amphimic-
tic. The taxon described above accordingly must be regarded as a species.
If specific distinctness from Rosa’s f. “typica” is demonstrable, Michael-
sen’s name probably can be retained. At least it has priority over remain-
ing unplaced varietal names.

Further discussion of most relationships should be postponed until
other varieties of veneta have been adequately characterized.

In America, E. zebra is easily distinguished from its congeners: From
E. foetida, by its thicker soma, greater number of segments, wider pairing
of setae, more posterior invariant section of the clitellum, more posterior
anterior margin of tubercula pubertatis, calciferous lamellae largest in xi
(rather than xii), a more posterior typhlosole termination (usually in
region of 110th-130th rather than 80th—98th segments), a more posterior
junction of extra-esophageal and dorsal trunks (in xii instead of ix—x),
absence of epididymis in male gonoducts, greater development of atrial
glands, presence of TP glands and of acinous, supraparietal GS glands.
Many of such characters were derogated or ignored by previous special-
ists.

From E. hortensis (cf. Gates, 1968b), by the larger soma, greater num-
ber of segments, more obvious restriction of pigment to transverse in-
trasegmental bands, invariant portion of the clitellum comprising xxviii-
xxxiii (rather than xxviii-xxxii), typhlosole termination usually in 110th—
130th (rather than 72’d—92’'d) segments, etc.

Enterion roseum Savigny, 1826, (common in America), according to
American and some European zoologists is in Eisenia. Other Europeans
refer it to Allolobophora. The species belongs in neither genus but deter-
mination of its proper position awaits further lumbricid revisions. Sav-
igny’s species is readily distinguished from all Eisenia spp. by presence of
calciferous sacs in x and by the U-shape of nephridial vesicles.

ADDENDUM

European lumbricids now domiciled in North America are: Alloloboph-
ora chlorotica (Savigny, 1826), A. limicola Michaelsen, 1890, A. longa

Earthworm revision 459

Ude, 1895, A. muldali Omodeo, 1956, A. trapezoides (Duges, 1828),
A. tuberculata Eisen, 1874, A. turgida Eisen, 1874, Dendrobaena mam-
malis (Savigny, 1826), D. octaedra { Savigny, 1826), D. rubida (Savigny,
1826), Eisenia foetida (Savigny, 1826), E. hortensis Michaelsen, 1890, E.
rosea (Savigny, 1826), E. zebra Michaelsen, 1902, Eiseniella tetraedra
(Savigny, 1826), Lumbricus castaneus (Savigny, 1826), L. festivus (Sav-
igny, 1826), L. rubellus Hoffmeister, 1843, L. terrestris, Linnaeus, 1758,
Octolasion cyaneum (Savigny, 1826), O. tyrtaeum (Savigny, 1826).

E. zebra, like three other species, has not been intercepted from earth
with plant shipments.

LITERATURE CITED

Gates, G. E. 1966. Requiem for megadrile Utopias. A contribution to-
ward the understanding of the earthworm fauna of North
America. Proc. Biol. Soc. Washington, 79: 239-254.

. 1967. On the earthworm fauna of the Great American Desert

and adjacent areas. Great Basin Nat., 27: 142-176.
1968a. On two American genera of the earthworm family
Lumbricidae. Jour. Nat. Hist. London, (in press).
1968b. Contributions to a revision of the Lumbricidae. III.
Eisenia hortensis (Michaelsen, 1890). Breviora, Mus. Comp.
Zool. No. 300: 1-12.

Gerarp, B. M. 1964. Lumbricidae. Linnean Soc. London, Synopses
British Fauna, No. 6. London. Pp. 58.

Omopeo, P. 1952a. Oligocheti della Turchia. Ann. Mus. Zool. Univ.
Napoli, 4(2), pp. 1-20.

. 1952b. Cariologia dei Lumbricidae. Caryologia, 4, pp. 173—

275.
1954. Alcuni lombrichi delle Alpi Vente e della coasta
orientale dell’Adriatico. Atti Mus. Sto. Nat. Trieste, 19(3):
121-135.

SmituH, F, 1924. The calciferous glands of Lumbricidae and Diplocardia.
Illinois Biol. Mons., 9(1): 1-76.

460 Proceedings of the Biological Society of Washington

Vol. 8. Mn 3 October 1969

PROCEEDINGS
OF THE

BIOLOGICAL SOCIETY OF WASHINGTON, JU! 2

THE MONOGENEAN PARASITIES OF AFRICAN FISHES.

X. TWO ADDITIONAL DACTYLOGYRUS SPECIES FROM
SOUTH AFRICAN BARBUS HOSTS}!

C. E. Price, E. S. McCLELLAN, A. DRUCKENMILLER
AND L, G. JACOBS
Department of Biology, Millersville State College,
Millersville, Pennsylvania

The first African Dactylogyrus species were reported by
Price and Gery (in press). At this time approximately 20 spe-
cies of this genus are known from various regions of the con-
tinent.

This study consists of an account of two additional species
of Dactylogyrus recovered from South African fishes. One of
these, D. myersi new species, was recovered from the gills of
Barbus trimaculatus Peters. The other, D. varicorhini Bychow-
sky (1957), was harbored by Barbus kimberleyensis. This latter
dactylogyrid was initially reported as a parasite of Varicorhinus
capoeta taken in the Soviet Union. Paperna (1961) recovered
this monogenean species from both Barbus canis and Varicor-
hinus damascinus in Israel, both cyprinid species native to that
country.

The occurrence of the same parasite species on different host
genera separated by thousands of miles is deemed of sufficient
importance to warrant discussion at a later point.

Materials and Methods: The authors extend their thanks to
R. McC. Pott, Professional Officer, Provincial Fisheries In-
stitute, Lydenburg, Republic of South Africa for donation of
branchial materials utilized in this study and for identification
of host species.

1 This study jointly supported by: (1) Department of Biology, Millersville State

College and (2) a research grant from the American Philosophical Society (#4956—
Penrose fund ).

36—Proc. Biot. Soc. WasH., Vou. 82, 1969 (461)

462 Proceedings of the Biological Society of Washington

Host specimens were captured by seine and/or gill nets.
Branchial materials were frozen and then preserved in 3.5 per-
cent formalin prior to shipment to the United States. Gills and
recovered parasites were then treated as prescribed by Price
(1966) and measurements made as recommended by Price and
McMahon (1967). Appropriate measurements and _illustra-
tions were made with the aid of a calibrated filar micrometer
ocular and a camera lucida, respectively. Anatomical terms
employed were those recommended by Hargis (1958) and by
Price and Arai (1967). Average measurements are given first,
followed by minimum and maximum values enclosed in pa-
rentheses. All measurements are expressed in microns.

Research on Monogenea is steadily increasing. These para-
sites are being described from the fishes of many countries
where monogenetic trematodes were previously unknown. A
check of available recent literature indicates that well over 400
species have been described within the past 10 years. This
figure becomes more meaningful when it is realized that less
than 1000 species of Monogenea were known in 1957.

The senior author foresees an increasing number of taxo-
nomic inconsistencies and other difficulties which could be
considered a natural consequence of working in a difficult area
of research. I firmly believe, however, that the situation would
be vastly improved if all authors would include whole mount
illustrations of new species.

In a letter some time back, Dr. W. J. Hargis, Jr. (Director,
Virginia Institute of Marine Science, Gloucester Point, Vir-
ginia), mentioned the “hook and anchor” methods of many
workers in Monogenea. After working with these parasites for
some time, I now fully realize the import of this. I would like
to recommend that future new species have no status unless
a whole mount for each is provided. As a former “hook and
anchor” worker, I now feel that merely depicting sclerotized
structures constitutes at best an inadequate approach to tax-
onomic studies.

Dactylogyrus myersi new species

Host: Barbus trimaculatus Peters; family Cyprinidae.

Locality: Pongolo River, Lydenburg, Republic of South Africa.
Location of parasite on host: Gill filaments.

Two new monogeneans 463

Number studied: 12.
Holotype: USNM Helm. Coll. No. 70561.
Paratype: USNM Helm. Coll. No. 70562.

Description: A dactylogyrid of moderate size, provided with a smooth
cuticle, length 323 (298-339); greatest body width 94 (86-102), near
midlength. Anterior cephalic lobes well-developed, lateral lobes vestigial.
Pharynx prominent, quite muscular and subspherical in outline (both
dorsal and ventral views). Two pairs of eyespots, all members about
equal in size. Head organs (either side) consist of four glandular struc-
tures connected by a common duct; duct terminates in larger pharyngeal
gland. Peduncle short and stout, with result that haptor is not well dif-
ferentiated from body proper (Fig. 1: whole mount).

One pair of anchors (dorsal) (Fig. 2). Each anchor composed of:
(1) a solid base equipped with well-defined deep and superficial roots,
(2) a solid shaft and (3) a solid point; shaft and point meet in a con-
tinuous arc. A perforation occurs through anchor near junction of shaft
and point. Anchors relatively long, ca. one-third as long as overall body
length, anchor length 107 (100-112); width of base 18 (16-21). An-
chor bases connected by a bar which is atypical for genus: ends are
sheet-like and partially encircle anchor bases, length 47 (43-52) (Figs.
3, 4).

Haptoral hooks 14 (seven pairs), similar in shape and size (Figs. 1,
5) and arranged five pairs ventral on haptor, two pairs dorsal (Mizelle
and Crane, 1964). Each hook composed of: (1) a solid elongate base,
(2) a solid shaft and (3) a sickle-shaped termination provided with an
opposable piece. Hooks range from 20 to 24 in length. The so-called
“additional” hooks of Dactylogyrus (Mizelle and Price, 1963) are con-
sidered not to be hooks at all, but to be vestiges of a ventral pair of an-
chors, disappearing as evolution progressed. These structures were not
observed in D. myersi.

Copulatory complex composed of a cirrus and a basally articulated ac-
cessory piece (Figs. 6, 7). Cirrus tubular, of a narrow diameter and
arranged in a coil of ca. 1.5 turns; diameter of coil 25 (22-28). Acces-
sory piece of unusual structure for genus; whereas most dactylogyrids pos-
sess accessory pieces with simple rami, those of the present form cross
over each other distally. Length of accessory piece 28 (25-31). Testis
postovarian, subspherical in outline and slightly smaller than ovary. Vas
deferens appears to loop over intestinal limb, but not observed with
certainty. Prostatic reservoir bipartite. Vagina not observed with cer-
tainty.

Intestine bifid, limbs simple and becoming confluent posteriorly. Vitel-
laria well-developed; co-extensive with intestinal crura.

Etymology: This species is named in honor of Dr. George S. Myers of
the Division of Systematic Biology of Stanford University, in appreciation
of the vast amount of ichthyological information he has furnished to the
senior author.

464 Proceedings of the Biological Society of Washington

100

Ficures 1-7. Dactylogyrus myersi new species. 1, Entire worm (ven-
tral view). 2. Anchor. 3, 4. Dorsal bar. 5. Hook. 6. Cirrus. 7. Acces-
sory piece.

Ficure 8. Anchor of D. varicorhini Bychowsky, 1957.

Discussion: Although Dactylogyrus can be considered a rather mor-
phologically homogeneous group, the present new species does not appear
to have any very close relatives. D. myersi possesses three characters
which are considered to be atypical for Dactylogyrus: (1) a bar with
modified ends which partially encircle the anchors, (2) relatively large
anchors with a perforation near the junction of shaft and point and (3)
an accessory piece in which one primary ramus crosses over the other.

Two new monogeneans 465

Dactylogyrus varicorhini Bychowsky, 1957

Host: Barbus kimberleyensis; family Cyprinidae.
Locality: Pongolo River, Lydenburg, Republic of South Africa.
Number studied: Twenty-six.

Previously Reported Hosts and Localities: (1) Varicorhinus capoeta,
in the Soviet Union, by Bychowsky (1957), (2) Barbis canis, in Israel, by
Paperna (1961) and Varicorhinus damascinus, in Israel, by Paperna
(1961).

Discussion: This species of Dactylogyrus is readily identified by ref-
erence to the anchors, which differ appreciably from those of the other
approximately 375 species of this genus. An anchor is depicted in Fig. 8.

The specimens in our possession agree quite well with the morphologi-
cal descriptions of D. varicorhini furnished by Bychowsky (1957) and
Paperna (1961). In size, our specimens are intermediate between those
described by these authors.

As noted above, D. varicorhini has been recovered from species of the
cyprinid genera Varicorhinus (Soviet Union and Israel) and Barbus
(Israel and South Africa). It is interesting to note that a given species of
parasite occurs on different host genera which are separated by thou-
sands of miles.

In a similar situation (Price and Yurkiewicz, in press) several specimens
of the monogenean genus Dogielius Bychowsky (1936) were recovered
from host specimens belonging to the genus Labeo in South Africa. The
original report by Bychowsky concerned the host genus Schizothorax. The
African and Soviet forms were separated by thousands of miles, as in
the case of Dactylogyrus varicorhini. Paperna (1961) reported Dogielius
from Varicorhinus in Israel.

One possible explanation for the occurrence of specific parasites on
widely separated hosts involves early stages in the evolution of cyprinid
fishes. Many ichthyologists believe that the cyprinids (family Cyprini-
dae) had their origin in Asia (Norman and Greenwood, 1963; Lagler,
Bardach and Miller, 1962). These fishes likely evolved from a characoid
ancestor. As Myers (1967) put it: “Cyprinoid fishes evolved in Asia from
some toothless characoid which got across the Tethys from Africa. In Eu-
rasia, the cyprinids blossomed into the largest familial group of the Ostario-
physi, and in the Tertiary invaded both Africa (across the greatly
shrunken Tethys) and North America (via a Bering land bridge).”

As the cyprinids underwent a veritable explosion of speciation, certain
of them apparently migrated toward Europe and the Northwestern part
of the Soviet Union. Others, as Myers (op. cit.) pointed out, crossed
what remained of the Tethys Sea into Africa. It is conceivable that an
ancestral minnow gave rise to two similar groups; one group headed
northwest, the other southwest. The ancestral form was likely a Barbus-
like form; offspring gave rise to Barbus as we know the genus today. This
genus maintained its identity and also gave rise to both Schizothorax and
Varicorhinus.

466 Proceedings of the Biological Society of Washington

Parasitological inference is that the genera Schizothorax, Varicorhinus
and Barbus are quite closely related. Paperna (1961) concurs in this.
Crass (1964) places some doubt upon the validity of Varicorhinus, believ-
ing that the genus might well be synonymous with Barbus.

If it is accepted that the cyprinid genera above are very closely related,
there remains only the necessity of accepting the well-established tenet of
parallel evolution that similar hosts harbor similar parasites to theorti-
cally account for the wide-spread occurrence of the parasites discussed
here.

LITERATURE CITED

BycHuowsky, B. E. 1936. Die Monogeneitschen Trematoden der Fische
des Tschu-Flusses. Trav. Exped. et Republ. Kirghiz, Moscow.
3: 245-275.

1957. Studies on Monogenoidea from Tadjikistan fishes.
(Russian text, with German summary). Isv. Vsiess. Nauchn.
Issl. Inst. Osior i Riechn. Rybn. Khos. 42: 109-123.

Crass, R. S. 1964. Freshwater fishes of Natal. Shutter and Shooter.
Pietermaritzburg, South Africa. 167 p.

Harcis, W. J., JR. 1958. A revised, annotated list of terms useful for
morphological studies of monogenetic trematodes. (Mimeo-
graphed at Virginia Marine Laboratory, Gloucester Point,
Virginia. 12 pp.)

Lacuer, K. F., J. E. BARpAcH, AND R. R. Miter. 1962. Ichthyology.
John Wiley and Sons, Inc. New York. 545 p.

MIzELLE, J. D., AND J. W. Crane. 1964. Studies on monogenetic trema-
todes. XXIII. Gill parasites of Micropterus salmoides (La-
cépéde) from a California pond. Trans. Amer. Microscop.
Soc. 83: 343-348.

, and C. E. Price. 1963. Additional haptoral hooks in the
genus Dactylogyrus. J. Parasitol., 49(6): 1028-1029.

Myers, G. S. 1967. Zoological evidence of the age of the South Atlantic
Ocean. Studies in Tropical Oceanography. Miami, 5: 614—
621.

Norman, J. R., anp P. H. GreENwoop. 1963. A history of fishes. Hill
and Wang, New York. 398 p.

ParpernA, I. 1961. Studies on monogenetic trematodes in Israel. 3.
Monogenetic trematodes of the Cyprinidae and Claridae of
the Lake of Galilee. Bamidgeh, 13(1): 14-29.

Price, C. E. 1966. Urocleidus cavanaughi, a new monogenetic trematode
from the gills of the keyhole cichlid, Aequidens maroni (Stein-
dachner). Bull. Georgia Acad. Sci., 24: 117-120.

, AND H. P. Arar. 1967. A proposed system of anatomy for
freshwater Monogenea. Canadian J. Zool., 45(6): 1283-
1285.

, AND T. E. McCManon. 1967. The monogenetic trematodes of
North American freshwater fishes. Riv. Parassit., 28: 177—
220.

Two new monogeneans 467

, AND J. Gery. (In press). Parasites des Poissons du Gabon. I.
Generalites sur les Trematodes monogenetiques, et descrip-
tion de six nouvelles especes parasites du genre Barbus. Bio-
logica Gabonica.

, AND W. J. YurkKiEwicz. (In press). The monogenean para-
sites of African fishes. VIII. A re-evaluation of the genus
Dogielius Bychowsky, 1936, with the description of a new
species. Rev. Iberica Parasitol.

468 Proceedings of the Biological Society of Washington

Vol. hi 3 October 1969

PROCEEDINGS
OF THE

ene

\\

BIOLOGICAL SOCIETY OF WASHINGTON \~

TAXONOMIC STATUS OF THE SHREW, NOTIOSOREX

(XENOSOREX ) PHILLIPSII SCHALDACH, 1966
(MAMMALIA: INSECTIVORA)

By Jerry R. CHOATE
Museum of Natural History, The University of Kansas,
Lawrence, Kansas

Among 129 mammals collected in southern Oaxaca in 1964
by Allan R. Phillips and William J. Schaldach, Jr., were four
short-tailed shrews, all tentatively identified as Cryptotis mex-
icana (Coues). Schaldach later discovered that two of the
specimens had only three “unicuspids” in each upper toothrow
instead of four (the normal complement for Recent species of
the genus Cryptotis Pomel). Further examination convinced
him that three (one lacking skull) of the four specimens rep-
resented an undescribed taxon; he assigned the fourth to
Cryptotis mexicana machetes (Merriam ).

The only Recent New World shrews that normally have but
three “unicuspids” in each upper toothrow are representatives
of the genera Notiosorex Coues and Megasorex Hibbard, which
many authors consider as congeneric (Notiosorex having prior-
ity). Although Schaldach (1966: 289-290) questioned the
“natural validity” of dental formulae as criteria for generic
determinations of shrews, he apparently failed to consider the
possibility that his specimens might represent a genus normally
characterized by the presence of more than three upper “uni-
cuspids.” Instead, he relied entirely on the dental formula for
generic allocation and (op. cit.: 289) named and described
Notiosorex phillipsii, setting it off in a separate subgenus
(Xenosorex ) characterized by its close resemblance to Crypto-
tis in characters other than number of teeth.

In his review of the Soricidae, Repenning (1967) placed

37—Proc. Biot. Soc. WAsH., VoL. 82, 1969 (469 )

>

470 Proceedings of the Biological Society of Washington

Notiosorex and Cryptotis in separate tribes (Neomyini and
Blarinini, respectively ) representing phylogenetic lineages that
probably have been distinct since early Miocene time (op. cit.:
61). This naturally aroused questions as to the identity and
status of Notiosorex phillipsii. Furthermore, my examination
of the holotype and paratypes of N. phillipsii revealed that on
the basis of external characters they cannot be distinguished
from the specimen assigned to C. mexicana caught at the same
locality, and that cranially the specimen of mexicana and the
two phillipsii accompanied by skulls differ only in the presence
or absence of the minute fourth upper “unicuspid.”

To determine the correct generic identity of phillipsii, the
one paratype (KU 114226) and the notes taken on the holotype
(UNAM 8445) and the other paratype (UNAM 5447) were
compared with representatives of each of N. crawfordi (Coues )
and N. evotis (Coues), the two nominal species of Notiosorex,
with Megasorex gigas (Merriam), and with representatives of
four species of Cryptotis—C. pergracilis nayaritensis Jackson,
C. mexicana mexicana (Coues), C. goodwini Jackson, and C.
magna (Merriam). The four species of Cryptotis were chosen
as representatives of morphologically distinct lineages within
that genus. Characters used by Repenning (op. cit.) to dis-
tinguish the Blarinini (p. 37) and Neomyini (p. 45) were eval-
uated and then applied to the study of phillipsii. Characters
used in diagnoses of the genera Cryptotis (p. 39), Notiosorex
(p. 55), and Megasorex (p. 56) were treated in a like manner.
Osteological and dental terminology and most of the diagnostic
characters used herein are from Repenning (op. cit.), except
that diagnostic characters have been modified slightly where
necessary to encompass the range of variation in Recent taxa.
The characters discussed below were chosen as most demon-
strative of relationships.

Dental formula: In Cryptotis the dental formula is 1-5—3/1-2-3 in Re-
cent species and all known fossil species except C. adamsi (Hibbard), in
which it is 1-6—3/1-2-3. In Notiosorex and Megasorex the dental for-
mula is 1|-4—3/1—2-3, the same as in specimens of phillipsii.

Cingular structure of “unicuspids”: In Cryptotis a more-or-less distinctly
developed cingular cusp, usually pigmented, is situated on the posterior
end of the lingual cingulum of each anterior upper “unicuspid.” In Notio-

Taxonomic status of Notiosorex 471

WAR A
LAR

FicurE 1.—Dorsal outlines of skulls of (a) Megasorex gigas (99538),
(b) Notiosorex evotis (90581), (c) N. crawfordi (89210), (d) N. phil-
lipsii (114226), (e) Cryptotis mexicana mexicana (29533), (£) C. magna
(99539), (g) C. goodwini (64610), and (h) C. pergracilis nayaritensis
(105408) showing degree of development of zygomatic process of maxil-
lary. KU catalogue numbers (in parentheses) apply to respective draw-
ings in Figs. 1-4.

472 Proceedings of the Biological Society of Washington

Ficure 2.— Mandibular articulation in (a) Megasorex gigas, (b)
Notiosorex evotis, (c) N. crawfordi, (d) N. phillipsii, (e) Cryptotis mexi-
cana mexicana, (f) C. magna, (g) C. goodwini, and (h) C. pergracilis
nayaritensis.

sorex and Megasorex the entire lingual cingulum may be elevated, form-
ing a cingular ridge that never is pigmented. Pigmented cingular cusps
are present in phillipsii and are similar to those in the species of Cryptotis
examined.

Pigmentation of teeth: In Cryptotis all teeth except the fourth upper
“unicuspid” are pigmented, the degree of pigmentation varying in dif-
ferent taxa. In Notiosorex the tips of the paracone of P4, protoconid of
ml, and some of the more anteriorly-situated teeth are variably pigmented.
In Megasorex pigmentation is lacking or at best slight. In phillipsii the
tips of the teeth are pigmented as in Cryptotis.

Degree of development of zygomatic process of maxillary: In Crypto-
tis the zygomatic process of the maxillary extends posterior from a place

Taxonomic status of Notiosorex 473

f g h

Ficure 3.—Structure of internal temporal fossa in (a) Megasorex gigas,
(b) Notiosorex evotis, (c) N. crawfordi, (d) N. phillipsii, (e) Cryptotis
mexicana mexicana, (f) C. magna, (g) C. goodwini, and (h) C. per-
gracilis nayaritensis. Note, as in other figures, the similarity between
phillipsii and C. mexicana.

opposite the metacone or metastyle of M2 as a short but distinct process
from which the masseter muscle originates. In Notiosorex the process
originates opposite the metastyle of M2 and either does not extend poste-
riad (N. crawfordi) or does so only as a minute process that probably

474 Proceedings of the Biological Society of Washington

f g h

Ficure 4.—Location of external temporal fossa in (a) Megasorex gigas,
(b) Notiosorex evotis, (c) N. crawfordi, (d) N. phillipsii, (e) Cryptotis
mexicana mexicana, (f) C. magna, (g) C. goodwini, and (h) C. per-
gracilis nayaritensis.

lacks significant muscular attachment (N. evotis). In Megasorex the proc-
ess originates posterior to M2 and does not extend posteriad. In _ phil-
lipsii the zygomatic process of the maxillary originates and extends poste-
riorly as in Cryptotis (Fig. 1).

Mandibular articulation: In Cryptotis the lingual condylar emargina-
tion is at least partially (usually considerably) filled with bone, varying

Taxonomic status of Notiosorex 475

in different species, so that the interarticular area is broad. In Notiosorex
and Megasorex the lingual condylar emargination is not filled, resulting in
a narrow interarticular area; the lower condyle is offset lingually (more
so than in Cryptotis) from the lower sigmoid notch, and is usually sep-
arated from that notch by a small groove. In phillipsii the mandibular
articulation is identical with that of Cryptotis (Fig. 2).

Structure of internal temporal fossa: In Cryptotis the internal temporal
fossa tends to be large, triangular, and excavated dorsally in such a fash-
ion that a basin is formed above the fossa proper. In Notiosorex and Mega-
sorex the fossa tends to be small, deep, and round, lacking all but a hint
of excavation. The structure of the internal temporal fossa in phillipsii is
identical with the condition found in Cryptotis (Fig. 3).

Location of external temporal fossa: In all species examined of Cryp-
totis the external temporal fossa is situated high on the coronoid process,
extending down no farther than the superior sigmoid notch. In Notiosorex
and Megasorex the fossa is situated low on the coronoid process, the ven-
tral margin often extending as low as the lower articular facet. In phil-
lipsii the fossa is situated as in Cryptotis (Fig. 4).

As shown above, specimens referred to “Notiosorex (Xenosorex) phil-
lipsiv’ clearly share morphological affinities, excepting dental formula,
with Cryptotis rather than Notiosorex. Examination of the specimens of
Cryptotis mexicana mentioned above and of additional material (ENCB
3413-14; AMNH 213758-59, 214152, 214803-06, 214808-09; UMMZ
112572) from near the type locality of phillipsii demonstrated that the
fourth upper “unicuspid” is variable in size and development in that
population, and that absence of the tooth does not constitute a valid
taxonomic character even at the subspecific level. Therefore, Xenosorex
hereby is transferred to the genus Cryptotis (in which it becomes an
available junior synonym), and phillipsii is placed in the synonymy of
Cryptotis mexicana peregrina (Merriam). The complexities of specific
allocation of the nominal subspecies of C. mexicana is beyond the scope of
the present paper, but will be discussed in a forthcoming review of Mid-
dle American shrews of the genus Cryptotis.

ACKNOWLEDGMENTS

I am grateful to J. Knox Jones, Jr., of The University of Kan-
sas Museum of Natural History (KU), Bernardo Villa R. of
the Universidad Nacional Autonoma de México (UNAM),
Ticul Alvarez S. of the Escuela Nacional de Ciencias Biologias
(ENCB), Richard G. Van Gelder of the American Museum of
Natural History (AMNH), and William H. Burt of The Uni-
versity of Michigan Museum of Zoology (UMMZ) for permis-
sion to examine specimens. Carleton J. Phillips and Hugh H.
Genoways kindly reviewed the manuscript, and Guy G. Musser
and Ticul Alvarez provided helpful comments and suggestions.

476 Proceedings of the Biological Society of Washington

Illustrations were prepared by Rosemary Fidelis Choate. Funds
enabling the author to examine specimens in México were pro-
vided by a Watkins Museum of Natural History Grant, The
University of Kansas, and funds for travel to other museums
listed above were made available in the form of a grant from
the National Science Foundation through the Committee on
Systematics and Evolutionary Biology at The University of
Kansas.
LITERATURE CITED
REPENNING, C. A. 1967. Subfamilies and genera of the Soricidae. .. .
U.S. Geol. Surv. Prof. Paper, 565: iv + 74.

ScHALDACH, W. J., Jn. 1966. New forms of mammals from southern

Oaxaca, Mexico, with notes on some mammals of the coastal
range. Saugetierk. Mitt., 4: 286-297.

Vol. 82, pp. 477-488 17 November 1969

PROCEEDINGS
OF THE

BIOLOGICAL SOCIETY OF WASHINGTON

A NEW PUFFER FISH, SPHOEROIDES PARVUS, FROM
THE WESTERN GULF OF MEXICO, WITH A KEY TO
SPECIES OF SPHOEROIDES FROM THE ATLANTIC

AND GULF COASTS OF THE UNITED STATES

By Ropert L. Surpp AND RALPH W. YERGER
Department of Biological Science
Florida State University, Tallahassee, Florida

Our taxonomic studies of the puffers (family Tetraodonti-
dae) in the Atlantic Ocean and adjacent waters have revealed
that the dominant inshore representative of the genus Sphoeroi-
des Anonymous in the western Gulf of Mexico is an un-
described and endemic species. This discovery is especially rel-
evant to the controversy concerning the relationship of the fish
faunas of the eastern and western Gulf of Mexico. Baughman
(1950: 118), Ginsburg (1952: 101), and Briggs (1958: 244)
considered the faunas to be relatively distinct, and the latter
author cited ecological evidence from Hedgpeth (1954: 206) to
justify this view. Hildebrand (1954: 232) held the opposite
opinion, and pointed to the apparent lack of evidence for en-
demic forms in the western Gulf of Mexico.

We are grateful to the following persons and their institu-
tions (with abbreviations used in this paper) for loan of
material: James C. Tyler, Academy of Natural Sciences of
Philadelphia (ANSP); Frederick H. Berry, (formerly of) U.S.
Bureau of Commercial Fisheries, Brunswick, Georgia (BLBG);
Donald Moore, U. S. Bureau of Commercial Fisheries, Galves-
ton, Texas (BLGT); Charles E. Dawson, Gulf Coast Research
Laboratory (GCRL); Royal D. Suttkus, Tulane University
(TU); Victor G. Springer, Smithsonian Institution (USNM);
Herbert T. Boschung, University of Alabama (UA); Carter R.
Gilbert, University of Florida (UF); Henry H. Hildebrand,

38—Proc. Bro.. Soc. WasuH., Vou. 82, 1969 (477)

478 Proceedings of the Biological Society of Washington

Ficure 1. Sphoeroides parvus n. sp., A. holotype, 79.7 mm SL. Mobile
Bay, Alabama, 8 August 1967. B. paratype, FSU 15365, from same series
as holotype, 75.3 mm SL.

University of Corpus Christi, who supplied material from the
Institute of Marine Science, University of Texas (IMS); and C.
Richard Robins, University of Miami Marine Laboratory
(UMML). Additional material was from the Florida State
University (FSU) collection. Our especial thanks are ex-
tended to James R. Martin who aided in the collection of ma-
terial, and to Dr. Victor G. Springer who reviewed the manu-
script.

The terminology, counts, and measurements follow Hubbs
and Lagler (1958: 19-26) except for the modifications dis-
cussed by Shipp and Yerger (1969: 425).

Sphoeroides parvus new species
Least puffer
Fig. 1
Sphoeroides marmoratus. Gunter, 1945: 84.
Sphoeroides nephelus. Hildebrand, 1954: 320. Hildebrand, 1955:
218 (in part). Reid, 1955: 331. Hoese, 1958: 347. Miller, 1965:
103.
Sphaeroides nephelus. McFarland, 1963: 100. Parker, 1965: 218.
Holotype: USNM 203248, an adult female 79.7 mm standard length
(SL), collected in a shrimp trawl near the center of Mobile Bay, Alabama,
by R. L. Shipp and J. Martin, 8 August 1967.
Paratypes: Twenty-five series comprising 382 specimens from the

northern and western Gulf of Mexico. FLORDIA: UMML 2618 (1

A new puffer fish 479

specimen, 62 mm SL), Apalachicola, 10 October 1950. UF 4437 (8, 54—
69), Choctawhatchee Bay, East Pass, 11 December 1954. UF 2731 (2,
52-57), Pensacola, 3.5 mi. E of Inerarity Pt., 15 August 1953. BLBG (1,
53), lower Pensacola Bay, between ship channel and south shore from
Big Lagoon to USCG station, 20 February 1964. ALABAMA: UA 62
(5, 47-64), Gulf Shores, 29 April 1950. FSU 15364 (35, 32-54), Mo-
bile Bay, 8 August 1967. FSU 15365 (176, 21-90), taken with holo-
type. UA 296 (15, 48-86), Mississippi Sound, 15 November 1952. UA
397 (5, 46-97), Mississippi Sound, 5 December 1953. UA 1290 (17,
47-79), Mississippi Sound, 18 April 1964. MISSISSIPPI: UA 625 (22,
30-58), Mississippi Sound, 18 October 1957. LOUISIANA: TU 9381
(1, 51), Lake Pontchartrain, 2 mi. W of South Draw, 30°10'N, 89°55’W,
5 November 1954. TU 22573 (14, 41-67), Gulf of Mexico, off Grand
Terre, 12 December 1959. ANSP 97647 (51, 42-75), Barataria Bay, 24
November 1931. TU 19038 (2, 80-90), Cameron, W bank Calcasieu
River, 28 April 1957. TEXAS: BLGT Gus 1 E25 (1, 56), 29°10’N,
89°42'W, January 1963. BLGT Gus 4 W1 (3, 58-62), 29°OI1'N, 95°
05’W, 2-7 March 1963. BLGT Gus 3 W138 (1, 62), 28°19’N, 96°21'’W,
1-6 April 1963. ANSP 98279 (Oregon station 3829) (2, 54-56), 28°
17.5’N, 93°57.5' W, 16 September 1962. BLGT Gus 1 W11 (2, 51-87),
27°42'N, 97°05'W, 2-5 February 1963. ANSP 98275 (2, 50-55), 26°18’
N, 97°11’W, September 1962. IMS 624 (1, 118), Aransas Bay, July
1956. MEXICO: IMS 614 (3, 62-71), off Pta. Frontera, 29 July—6
August 1951. IMS 619 (8, 63-75), Campeche to Champoton, 10-16
February 1951. IMS 622 (4, 80-85), W of Campeche, 27-29 July 1951.

Ten paratypes from FSU 15365 (Alabama, see above) have been sent
to each of the following institutions and assigned the indicated museum
number: American Museum of Natural History, AMNH 27399; Field
Natural History Museum, FNHM 74783; and Museum of Comparative
Zoology, MCZ 46203.

Other specimens: ALABAMA: UA 286 (1, 104), Mobile Bay, 11
October 1952. MISSISSIPPI: GCRL V65: 1284 (1, 51), S of Horn
Island, 28 August 1959. LOUISIANA: USNM 155990 (1, 70), Breton
Island, 12 March 1931. GCRL V66: 311 (1, 51), S of Grand Isle, 23
October 1958. TEXAS: USNM 156492 (3, 63-86), Freeport, Texas,
January—May, 1947. USNM 118648 (1, 88), Aransas Pass, 8 July 1941.
USNM 155989 (1, 111), Aransas Pass, 4 April 1929. USNM 155992 (1,
111), Corpus Christi Bay, 11 November 1926. USNM 73580 (1, 64),
Corpus Christi, 29 November 1891. ANSP 98263 (1, 71), Pt. Isabel, 30
November 1947.

Diagnosis: One of the smallest puffers (rarely exceeds 100 mm SL),
distinguished from other members of the genus by a combination of
characters: absence of lappets on dorsal surface of body, absence of
deeply pigmented spot in pectoral fin axil, snout short, interorbital re-
gion broad, flat (least bony interorbital width 25% or more of snout
length), and integument heavily covered by prickles, which do not extend
posteriorly beyond level of anus,

480 Proceedings of the Biological Society of Washington

E 10
= *S. parvus
= °S. nephelus ze ane
sy ee &
2 8 ec oe
= o 2? @2 ee 2 s
3 O) eferc> erely ane) ae |
3s ee eee re
-s e .
= 6 ° eo ad bd e
eo oe ° e
< Pe = owes °
— e °
oe .

a 4 % oe wo &
°
2
Pm
i |
o 2)
2 |

_ a= 8 12 16 20 24 28 32 36 40 44 48

Snout length (mm)

Ficure 2. Relationship between least bony interorbital width and
snout length in Sphoeroides nephelus and S. parvus. (Data on S.
nephelus from Shipp and Yerger, 1969).

Superficially, Sphoeroides parvus most closely resembles S. nephelus
(Goode and Bean) and S. maculatus (Bloch and Schneider). It differs
from both species by the lack of a deeply pigmented spot at the axil of
the pectoral fin. It further differs from S. nephelus in having a broad flat
interorbital region, a shorter snout [interorbital width less than one-
fourth snout length (Fig. 2)], and an irregular placement of the ventral-
most lateral spots (in S. nephelus these are arranged in an even row along
the ventrolateral body angle, see Fig. 3). It differs from S. maculatus
(Fig. 5D) in having the shape of the ventrolateral markings chiefly
round rather than vertically elongate, the prickles on the ventral surface
not extending beyond the anus, and by the absence of tiny jet-black
specks over most of the pigmented body surface.

Description: Body size small, snout short (18 percent SL), inter-
orbital region flat to slightly concave, very broad (5 percent SL). Anterior
body surface covered with close-set prickles or dermal spines, exposed in
both uninflated and inflated specimens; dorsally, prickles extend posteri-
orly to dorsal fin origin, and ventrally almost to anus.

Morphometric data for the holotype and 20 paratypes are given in
Table 1.

Fin ray counts of 50 type specimens chosen at random from through-
out the range of the species are as follows (the value including the holo-
type is italicized): dorsal rays 8 (in 43 specimens), 9 (7); anal rays 6
(4), 7 (45), 8 (1); pectoral rays (both sides counted separately) 13
(1), 14 (33), 15 (62), 16 (4).

Coloration: Ground color on dorsal surface brown or grey with scat-
tered, indistinct blotches or spots; laterally ground color fades slightly
above ventrolateral body angle; lower sides and ventral surface unpig-

A new puffer fish 481

dATP PEEP
O q 2 3 4 5 6 A 8 S

Ficure 3. A. Sphoeroides nephelus, FSU 15606, 182 mm SL, Key
Largo, Monroe Co., Florida, 29 December 1967. B. S. parvus, paratype,
FSU 15365, 75.3 mm SL, Mobile Bay, Alabama, 8 August 1967. Note in-
terspecific differences: size of adults, arrangement of lateral spots, and
snout length.

TABLE 1. Measurements of the holotype and 20 paratypes of Sphoeroi-
des parvus expressed in percent of standard length.

Paratypes*

Holotype Range Mean
Standard length 79.7 56-97 (Pal
Head length 36.1 33.4-38.5 Siar
Snout length 17.9 16.6-19.4 17.8
Least bony interorbital width Dio 4,4-6.3 Dell
Pectoral fin length 15.9 I5.0=2172 17.8
Depressed dorsal fin length L7.9 16.0—20.1 18.3
Depressed anal fin length Ibs 12782, 15.4
Caudal fin length 20.2 18.7-23.4 20.9
Snout to dorsal origin 71.4 66.0-74.8 70.5

*FSU 15365 (4 specimens); UA 397 (4); ANSP 97647 (2); TU 19038 (2),
22573 (1); BLGT: Gus 1 E25 (1), Gus 1 W4 (3), Gus 1 W111 (2), Gus 3
N 10k

482 Proceedings of the Biological Society of Washington

100, 90 = 80 : 70 a 60 |
=
ra 430

@ S. nephelus

é wr “Epa: 6) 4S. parvus
ete GDS 5S. parvus |
At

| ee ee
la a roe %
| ( Chek
| \ a X
| f | ae Cee
‘ a Bo aa ~ at 3 ite
* | ce (eee a ° |
) —s ee ee -
ee a : cb pet ee
ow = =

Ficure 4. General distribution of two species of Sphoeroides based on
specimens examined. Distribution of S. nephelus is included (based on
Shipp and Yerger, 1969) to indicate distributional patterns and zone of
sympatry with S. parvus.

mented. Lateral blotches or spots slightly more distinct than those on
dorsum; not always arranged in an even row, but tend to border ven-
tral boundary of ground color. Spot present in pectoral fin axil in few
specimens, but rarely more intensely pigmented than others on body. In-
distinct dark bar present between eyes. Dorsal and lateral surfaces often
with vague white specks, which may appear bright green in live speci-
mens. A few black specks on cheeks in some larger specimens. No other
noteworthy color marks appear in live specimens except for yellow or gold
cast over much of lateral and ventral surfaces. All fins unpigmented ex-
cept caudal, which may have an indistinct pigmented area near its base
and another near its distal end.

Adult size: S. parvus is the smallest known species of Sphoeroides in
the Atlantic Ocean and adjacent waters; the largest specimen examined
was 118 mm SL. Several authors previously noted the small size of this
puffer in the western Gulf (Gunter, 1945: 84; Hildebrand, 1954: 320;
Reid, 1955: 449; Miller, 1965: 103). Hildebrand (1955: 218) reported
a 91 mm (total length) female with nearly ripe ovaries. We have ex-
amined females as small as 55 mm SL and males 47 mm SL which were
sexually mature. The closely related species on the Gulf and Atlantic
coasts do not mature until a much larger size is attained (about 70 mm
SL in S. maculatus, usually more than 100 mm SL in S. nephelus), and
commonly exceed 150 mm SL (Fig. 3).

Distribution: S. parvus occurs from Apalachicola Bay, Florida, west-
ward throughout the western Gulf of Mexico. S. nephelus is the dominant
form in the clear waters of northwest Florida to Pensacola, but S$. parvus
replaces it in the muddy waters of Mobile Bay and westward (Fig. 4).
The senior author has examined many hundreds of puffers captured by
shrimp boats in Mobile Bay, and not one S. nephelus was found. This is

A new puffer fish 483

further verified by personal communication with the shrimpers who trawl
both the clear and muddy localities. Specimens of S. nephelus west of
Florida are rare. In the southwestern Gulf of Mexico, Hildebrand (1955:
218) reported both forms from the Campeche shrimp grounds, but S.
parvus was much more abundant.

Zoogeography: Sphoeroides parvus, S. maculatus, and S. nephelus con-
stitute a closely related species complex. Sphoeroides parvus, found in the
northern and western Gulf, is more closely allied morphologically to S.
maculatus, which occurs in the Atlantic from Canada to northeastern
Florida (Shipp and Yerger, 1969: 426), than to S. nephelus, a predomi-
nantly West Indian species which occurs on both coasts of Florida. We
believe that prior to the existence of the Florida peninsula, a continuous
population of puffers (the progenitor of S. maculatus and S. parvus) was
found around the southern coast of the United States. The emergence of
this peninsula split the population into two, one isolated in the Atlantic,
the other in the Gulf. Simultaneously the projection of this peninsula into
the tropical waters of the Caribbean provided suitable habitat for the
northward dispersal of S. nephelus from West Indian stocks. One or both
of these factors, a land barrier and competition with a closely related
species, apparently has maintained the isolation between the two original
coastal populations, and speciation has ensued. Meanwhile S. nephelus is
probably prevented from further dispersal northward and westward by
ecological barriers. The distribution of S. nephelus and S. parvus in the
Gulf of Mexico closely matches the ecologically distinct habitats described
by Hedgpeth (1954: 206).

This hypothesis supports that proposed by Springer (1959) for an ex-
planation of the strikingly similar distribution of the blenniid fishes,
Chasmodes bosquianus and C. saburrae, although the geologic dates
which he suggested for various shorelines may be erroneous.

Among other species of fishes with distributional patterns similar to
S. parvus in the northern and western Gulf are the sole, Gymnachirus
texae (see Dawson, 1964), the sparid, Stenotomus caprinus (see Cald-
well, 1955), and the cyprinodont, Fundulus confluentus pulvereus (see
Relyea, 1965).

KEY TO SPECIES OF SPHOEROIDES ON THE ATLANTIC AND
GULF COASTS OF THE UNITED STATES

Although the status of the puffers in the Southern and Eastern Atlan-
tic Ocean and parts of the Caribbean has not yet been studied satisfacto-
rily, the species which occur on the shores of the United States ( Atlantic
and Gulf of Mexico) are now sufficiently well known to provide a key to
facilitate their identification.

1A. Lappets (small fleshy tabs) present on dorsum; either a single,
black pair on the dorsum about one-half the distance between
the posterior margins of the orbits and the dorsal fin origin,
or many tan lappets (most easily seen when specimens are im-

484 Proceedings of the Biological Society of Washington

1B.
2A.

3A.

3B.

4A.

4B.

5A.

<

A new puffer fish 485

mersed in water) scattered on the posteriolateral and dorsolat-
ETL MES UTE] COS gemma Sat tel eRe ewe ee ee
Wea StS ma DSer te pee ee ke
A single pair of black lappets present on the dorsum. Cheeks
often marbled. From one to five poorly defined dark blotches
present on the lateral body surface posterior to the pectoral fin
Mier eee cemiees S. dorsalis Longley, Marbled puffer. Widespread in
western Atlantic and adjacent waters, in relatively deep
water (10-50 fathoms). Fig. 5A.
Many tan lappets present on the posterior portions of the body,
usually concentrated near the ventrolateral body angle. No mar-
bled pattern on cheeks. Five to eight (usually six or seven)
sharply defined, rounded lateral spots posterior to the pectoral
fin bordering the ventrolateral body angle —
Ear Ura cance S. spengleri (Bloch), Bandtail puffer. Widespread in
the western Atlantic and adjacent waters, in shallow
water. Fig. 5B.
Body variously mottled, not uniformly pigmented. Caudal dusky,
sometimes with pigment concentrated at base and distal end,
giving an indistinct barred appearance. Least bony interorbital
width 8.5 percent or less of SL
Body uniformly pigmented, except usually a few scattered spots
on dorsal and lateral surfaces. Caudal dusky except for distal tips
which are usually lighter. Least bony interorbital width 9 per-
COXEN OTE) (abst 0 OVO) ta O} cet) Lee eee es ne Sr Oe ree eee
One or two distinct, white, interorbital bars, the posterior often
connected by a posterior perpendicular extension to a dorsal
pattern of coarse white arches and circular markings
SAR reheat S. testudineus (Linnaeus), Checkered puffer. Wide-
spread in Caribbean, southern Gulf of Mexico, and
warmer waters of the western Atlantic, in shallow
water. Fig. 5C.
One vague, dark interorbital bar. No dorsal pattern of coarse
white arches
Several (usually six-eight) distinct, vertically elongate bars pos-
terior to pectoral fins. Dorsal and lateral surfaces in mature speci-

Ficure 5. A. Sphoeroides dorsalis, ANSP 105185, 127 mm SL, Tobago.
B. Sphoeroides spengleri, ANSP 104555, 111 mm SL, Columbia.
Sphoeroides testudineus, FSU 11928, 86 mm SL, Jupiter Inlet, Florida.
D. Sphoeroides maculatus, UF 11773, 171 mm SL, Georgia. E. Sphoeroi-
des nephelus, UMML 1366, 197 mm SL, Cocoa, Florida. F. Sphoeroides
parvus FSU 15365, 84 mm SL, Mobile Bay, Alabama. G. Sphoeroides
pachygaster, BLBG, Silver Bay 2190, 132 mm SL, Atlantic Ocean, off
South Carolina.

C.

486 Proceedings of the Biological Society of Washington

mens (above 70 mm) covered with tiny (1-2 mm) jet-black
spots. Prickles on ventral surface extend posteriorly beyond the
anus, usually to the anal fin origin. Pectoral rays 15-17, usually
NG Se a a A 2
— §. maculatus (Bloch and Schneider), Northern puffer.
Western North Atlantic, from Newfoundland to
northeast Florida, usually in shallow water. Fig. 5D.
5B. Spots present posterior to pectoral fins. No tiny jet-black spots
on dorsal or lateral surfaces. Prickles present or absent, but when
present, do not extend beyond the anus. Pectoral rays usually
I4-or 15 (rarely 1i8>or 16) 222 ee 6
6A. Spot at axil of pectoral fin more intense than any other on body.
Bony interorbit usually concave; least bony width narrow, more
than 4 in snout. Adults commonly exceed 125 mm SL
ee oes _. §. nephelus (Goode and Bean), Southern puffer.
Caribbean, eastern Gulf of Mexico, and Atlantic coast
of Florida, in shallow water. Fig. 5E.
6B. Spot at axil of pectoral fin absent, or if present, rarely more in-
tense than other spots on body. Bony interorbit nearly flat; least
bony width broad, less than 4 in snout. Not known to reach 120
ie S. parvus Shipp and Yerger, Least puffer. Northern
and western Gulf of Mexico, in shallow water. Fig. 5F.
7A. Body smooth. Caudal short, more than 6 in SL
Pec these tha! S. pachygaster (Miller and Troschel), Blunthead puf-
fer. Most of western Atlantic, in relatively deep water
(30-100 fathoms). Fig. 5G.
7B. Prickles on dorsal and ventral surface. Caudal moderately long,
eloul-0 In, iy Gos 0 ee
i Sta Oe S. trichocephalus (Cope), Hairy puffer. Known from
one specimen washed ashore at Rhode Island. Not fig-
ured.
LITERATURE CITED
BAUGHMAN, J. L. 1950. Random notes on Texas fishes. Part I. Texas
Jour. Sci. 1: 117-138.
Briccs, J. C. 1958. A list of Florida fishes and their distribution. Bull.
Fla. State Mus. (Biol. Sci.) 2(8): 223-318.
CaLpwe.Li, D. K. 1955. Distribution of the longspined porgy, Stenoto-
mus caprinus. Bull. Mar. Sci. Gulf and Carib. 5(2): 230-
DRSKSY:
Dawson, C. E. 1964. A revision of the western Atlantic flatfish genus
Gymnachirus (the naked soles). Copeia 1964 (4): 646-665.
Gryspurc, I. 1952. Eight new fishes from the Gulf coast of United
States, with two new genera, and notes on geographic distri-
bution. Jour. Wash. Acad. Sci. 42(3): 84-101.
Gunter, G. 1945. Studies on marine fishes of Texas. Publ. Inst. Mar.
Sci. Univ. of Texas 1(1):; 1-190.

A new puffer fish 487

Hepcpetu, J. W. 1954. Bottom communities of the Gulf of Mexico. In:
P. S. Galtsoff (coordinator ), Gulf of Mexico, its origin, water,
and marine life. Fish. Bull. U. S. Fish and Wildl. Ser. 55
(89): 203-214.

Hintpesranp, H. H. 1954. A study of the fauna of the brown shrimp
(Penaeus aztecus Ives) grounds in the western Gulf of Mex-
ico. Publ. Inst. Mar. Sci. Univ. of Texas 3(2): 234-366.
1955. A study of the fauna of the pink shrimp grounds in
the Gulf of Campeche. Publ. Inst. Mar. Sci. Univ. of Texas
4(1): 169-232.

Horse, H. D. 1958. A partially annotated checklist of the marine fishes
of Texas. Publ. Inst. Mar. Sci. Univ. of Texas 5: 312-352.

Husps, C. L. anp K. L. LAGLER. 1958. Fishes of the Great Lakes region
(revised edition). Bull. Cranbrook Inst. Sci. 26: 1-186.

McFarvanp, W. N. 1963. Seasonal changes in the number and biomass
of fishes from the surf at Mustang Island, Texas. Publ. Inst.
Mar. Sci. Univ. of Texas 9: 91-105.

Miter, J. M. 1965. A trawl survey of the shallow gulf fishes near Port
Aransas, Texas. Publ. Inst. Mar. Sci. Univ. of Texas 10: 80-
107.

Parker, J. C. 1965. An annotated checklist of the fishes of the Galveston
Bay system, Texas. Publ. Inst. Mar. Sci. Univ. of Texas 10:
201-220.

Rem, G. K. 1955. A summer study of the biology and ecology of East
Bay, Texas. Part II. Texas Jour. Sci. 7(4): 430-453.

Retyea, K. G. 1965. Taxonomic studies of the cyprincdont fishes, Fun-
dulus confluentus Goode and Bean, and Fundulus pulvereus
(Evermann). M. S. thesis, Florida State Univ., Tallahassee,
Fla.

SHipp, R. L. AnD R. W. Yercer. (1969). Status, characters, and dis-
tribution of the northern and southern puffers of the genus
Sphoeroides. Copeia 1969 3: 425-433.

SPRINGER, V. G. 1959. Blenniid fishes of the genus Chasmodes. Texas
Jour. Sci. 11: 321-334.

488 Proceedings of the Biological Society of Washington

Vc aie 17 November 1969 .

PROCEEDINGS aa re ee
OF THE \ NUV oo l0e

eee AL SOCIETY OF WASHINGTON “\Ligparif2~

CHAPINIA ELBELI TENDEIRO, A SYNONYM OF
CHAPINIA FASCIATI ELBEL (MALLOPHAGA:
MENOPONIDAE )

By Rosert E. ELBEL
Ecology and Epidemiology Division, Deseret Test Center,
Dugway, Utah

Tendeiro (1967) described Chapinia elbeli from two males
and one female off Tockus alboterminatus stegmanni (Neu-
mann), but Elbel (1967) previously described C. fasciati from
T. f. fasciatus (Shaw) as type host with paratypes from T. a.
suahelicus (Neumann).

Through the courtesy of Dr. Tendeiro, the holotype, allotype
and paratype of C. elbeli were examined and appear to be
morphologically identical with C. fasciati from the type host,
from T. a. stegmanni in the American Museum of Natural His-
tory, and from T. a. australis (Roberts) in the United States
National Museum.

Tendeiro stated that C. elbeli differed from C. fasciati and
from other members of the lophocerus species group in the
male genitalia by the external indented swelling near the
posterior end of the parameres, by the two fingerlike posterior
points of each lateral horn, and in the female by the absence of
sclerital hooks on each side of the midline of the ventral scle-
rite between the vulva and anus. In addition he stated that
the female anal fringe had 62 setae. Although not mentioned or
illustrated by Elbel, the indented swellings on the parameres
are present in all members of the lophocerus species group, and
are the sockets from which the parameres are split posteriorly,
a character which separates the other two species groups from
the lophocerus. An examination of Tendeiro’s specimens shows
that both males do indeed possess Pye posterior points

39—Proc. Brox. Soc] Wasx., Vou. 82, 1969 (489)

i

490 Proceedings of the Biological Society of Washington

on each lateral horn of the genitalia as in C. fasciati, and the
female does have sclerital hooks on each side of the midline of
the ventral sclerite between the vulva and anus as in all mem-
bers of the lophocerus species group. However, the female
anal fringe has 64 setae in Tendeiro’s specimen, 66 and 68 in a
specimen each from the American Museum of Natural History
and the United States National Museum. Thus, the range of
the anal fringe of C. fasciati is 64-86 rather than 70-86 as given
by Elbel. C. camuri Elbel has an anal fringe of 60-64 setae; but
C. fasciati from both host species, including Tendeiro’s speci-
men, has the ventral sclerite between the vulva and anus ele-
vated medially between the sclerital hooks more than in C.
camuri.
LITERATURE CITED
ELBEL, Ropert E., 1967. Amblyceran Mallophaga (biting lice) found
on the Bucerotidae (Hornbills). Proc. U. S. Nat. Mus. 120:
1-76.
TENDEIRO, JoAo, 1967. Etudes sur les Mallophages. Mallophages du

Parc National de ’Upemba (Congo) (Mission G. F. de
Witte). Rev. Estud. Gerais Univ. Mocambique 4: 361-441.

[7s

~~

Vol. & 17 November 1969

PROCEEDINGS
OF THE

BIOLOGICAL SOCIETY OF WASHINGTON

NOTROPIS XANTHICARA, A NEW CYPRINID FISH
FROM THE CUATRO CIENEGAS BASIN, NORTH-
CENTRAL MEXICO

By W. L. MINCKLEY AND Gaby L, LYTLE
Department of Zoology, Arizona State University, Tempe 85281

The lutrensis-ornatus group of the subgenus Cyprinella of
Notropis (Gibbs, 1957) includes in the middle Rio Grande and
adjacent drainages a number of nominal and undescribed forms
related to Notropis proserpinus (Girard). Notropis proserpinus,
as understood by us, lives in clearer waters of the lower Pecos
River system from southern New Mexico (Koster, 1957) south
and east to the Devil’s River and San Felipe Spring, Val Verde
County, Texas. Koster’s implication (p. 68) that this species
is present in the lower Rio Grande Valley, New Mexico, is un-
verified. Notropis lepidus (Girard) inhabits spring-fed waters
of the Nueces, Frio, Medina, and Guadalupe River systems in
Texas (Hubbs, 1954). Notropis rutilus (Girard) is in similar
habitat in the rios Salado and San Juan, northern México. And,
two undescribed species, one from the basin of the Rio Con-
chos, Chihuahua, México (Salvador Contreras B. and Min-
ckley, unpublished ), and the other from the semi-isolated Bol-
son of Cuatro Ciénegas, central Coahuila, northern México
(Minckley, 1969), complete the assemblage. The present con-
tribution describes the Cuatro Ciénegas form, and is one in a
series of papers resulting from research supported by N.S.F.
Grants GB-2461 and GB-6477X. Thanks are due the curators
of various museums for aid in obtaining specimens, the person-
nel who assisted in collections, and persons who assisted in
analyses. Permission to collect in various states, and in Mexico,
readily granted, is gratefully acknowledged.

40—Proc. Biot. Soc. WaAsH., Vou. 82, 1969 (491)

492 Proceedings of the Biological Society of Washington

Ficure 1. Male and female paratypes of N. xanthicara (KU 7404;
upper) and a male and female of N. rutilus from the Rio Salado de los

Nadadores (KU 7347; lower).

New Mexican fish 493

Notropis xanthieara new species
(Notropis sp., Minckley, 1969)
Cuatro Ciénegas shiner (Fig. 1)

Diagnosis: Notropis xanthicara is a member of the subgenus Cyprinella
of Notropis, as delimited by Gibbs (1957), and is most closely related to
the nominal N. rutilus, from which it probably arose. The new species
may be distinguished from the latter, and from other members of the sub-
genus, by the following combination of characters: body terete, not ob-
viously slab-sided; lateral-line scales usually 34, often 33; anal fin-rays
usually eight; pharyngeal teeth 4—4, slender and hooked, with serrated
grinding edges; lower jaw included; lateral band black, discrete, about
one scale-row wide, extending to, but not through, eye, appearing again as
as a pre-ocular streak on each side of snout; scale pockets weakly outlined
on abdomen; belly immaculate, with black peritoneum showing through
midline; predorsal streak broad and diffuse; postdorsal streak faint to ab-
sent; gular area variably darkened, interopercular area and anterior part
of breast sometimes bearing melanophores; breeding male predominantly
yellow, especially on head and fins; nuptial tubercles on snout of breeding
males separated from those on dorsum of head by a distinct hiatus.

Material: About 1,000 specimens of Notropis xanthicara were ex-
amined, from throughout the Cuatro Ciénegas basin. Detailed locality
data are provided only for the holotype and for paratypes. Information
on non-type material of N. xanthicara, and on comparative material of
N. rutilus, is by river system only. Detailed reports on range and varia-
tion of the entire species-group are in preparation. The abbreviations for
depositories are as follows: ASU = Collection of Fishes, Arizona State
University, Tempe; KU = Museum of Natural History, University of
Kansas, Lawrence; UMMZ = University of Michigan Museum of Zool-
ogy, Ann Arbor; and UNL, Laboratorio de Vertebrados, Universidad de
Nuevo Léon, Monterrey. Additional materials, loaned by Tulane Univer-
sity and the University of Texas, will be reported later. All localities in
the Cuatro Ciénegas basin are given in kilometers (km) from the center
of Cuatro Ciénegas. Field data has been revised to correspond to the
detailed map published by Minckley (1969), and reproduced here in
simplified form as Figure 2.

Notropis xanthicara: Holotype—UMMZ 188782, a mature, tuberculate
male, 45 millimeters (mm) long, collected 6 April 1961 by R. R. Miller
and family, C. L. Hubbs, D. R. Tindall, and W. L. Minckley, Rio Puente
Colorado, 8.5 km south and 0.7 km west of Cuatro Ciénegas, Coahuila,
México. Paratypes—UMMZ 179834, 11 specimens, collected with the
holotype; ASU 969, 8 specimens, collected at the type locality, 10 June
1964. ASU 2316, 185 specimens, collected 25 December 1965, Rio
Churince, 14.7 km south and 7.0 km west. UMMZ 179202, 46 specimens,
collected 6 April 1961; UNL 703, 6 specimens, and UNL 709, 105 speci-
mens, collected 26 and 27 August 1964, respectively, all from Posos de
la Becerra, 11.4 km south and 6.7 km west. ASU 3728, 68 specimens,

494 Proceedings of the Biological Society of Washington

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SIERRA DE MENCHACA Lee) fe
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Ficure 2. Distributional records for N. xanthicara in the Cuatro Ciéne-
gas basin, Coahuila, México; dots = localities for specimens in museums,
open circles = field observation of the species, and the circle dot = the
type locality (map modified from Minckley, 1969).

collected 16 August 1968, Rio Garabatal, 6.7 km south and 8.8 km west.
KU 7404, 72 specimens, collected 17 Apri! 1963, Canal de Julio, 3.4 km
south and 1.2 km east. UMMZ 179860, 53 specimens, collected 9 April
1961, Rio Mesquites, 8.2 km south and 0.8 km west. Additional ma-
terial (non-types )—Rio Churince system (see Minckley, 1969, for de-
tails on intra-basin drainages; numbers of specimens are in parentheses ) :
ASU 1658, 1747, 2332; KU 7389; UMMZ 179878 (99 fish). Posos de la
Becerra-Rio Garabatal system: KU 7374, 7383; UMMZ 179827 (56).
Lagunas de Juan Santos: UNL 714 (45). Rio Mesquites system: ASU
2268; KU 7363, 7394 (151). Rio Puente Chiquito system: UMMZ
179205 (125). Canal de Saca del Fuente: ASU 945 (2). Tio Candido
system: UMMZ 179221 (1). Santa Tecla system: KU 7421 (5).

Notropis rutilus: E. G. Marsh collection (discussed later): UMMZ
130377, 130387. Rio Salado system: ASU 913, 1723; KU 7347; UMMZ
130366, 130369, 130398, 179805, 179812; UNL 439, 688, 697. Rio San
Juan system: UMMZ 97420-7 (97421 = topotypes), 124425, 146980,
162131; UNL 29, 648.

Description and Comparisons: Notropis xanthicara is similar in most
respects to N. rutilus, differing principally in features of pigmentation and

New Mexican fish 495

in its slender, more delicate structure (Fig. 1). Fin-rays of the new form
are especially fragile. Anal rays, counted as the last two sharing a com-
mon base, number eight in 264 of 283 fish, ranging from seven (two fish )
through nine (17 fish). A similarly strong mode of eight anal rays is pres-
ent in 447 counts available for N. rutilus (six rays in one; seven in five;
eight, 371; and nine, 70).

Length of the dorsal fin in N. xanthicara is usually equal to or greater
than depth of body. In rutilus the length of the dorsal fin is most often
less than greatest depth of body. Shape of the dorsal is similar in both
species: the fin is square to slightly concave at its distal margin, and the
anterior rays reach approximately to the tip of the last, when the fin is
depressed. The anal fin of xanthicara tends to be slightly falcate, especially
in larger adults, with the first rays extending past the tip of the last when
depressed. In rutilus, the distal border of the anal fin is squared to
slightly falcate, but the first ray usually falls short of the tip of the last
(except in some breeding males). Caudal fins are large in both forms,
but in xanthicara the upper lobe is slightly more expansive, and the lower
more elongate and acutely tipped.

Standard lengths range from 30.2 to 56.0 mm in N. xanthicara (149
fish) and from 30.0 to 60.0 mm in our sample of rutilus (447 fish). Ori-
gin of the dorsal fin is slightly more posterior in N. xanthicara. Ranges
which follow mean values represent, first, the over-all range obtained,
and second, the range of means for 13 samples of xanthicara and for 23
populations of rutilus. Predorsal length averages 54.1 per cent of stan-
dard length (49.5-59.5; 53.1-55.6) in xanthicara, and is almost identical
in rutilus, 53.3 per cent (50.0-58.0; 51.2-55.7). Dorsal-fin origin is
behind the insertion of the pelvic fins in both species. Prepelvic lengths
are more different than predorsal lengths, averaging 52.2 per cent of
standard length for xanthicara (48.6—56.3; 50.3-53.7), and 50.2 per cent
for rutilus (46.9-55.0; 48.8—51.7).

The paired fins are relatively small in N. xanthicara. The pectoral fins
are acutely tipped, then gently rounded on their distal margins. They ex-
tend 2/3 to 3/4 of the distance to the pelvic-fin bases, and are slightly
longer in males. Pectoral fins of rutilus from the Rio Salado system are
similar, as are those of non-breeding specimens from the Rio San Juan basin.
However, some samples of males of rutilus from San Juan have expan-
sive, elongate pectoral fins. The pelvic fins of xanthicara are pointed,
with an almost-straight distal margin. Pelvic fins of rutilus are more
rounded and often longer.

Mean numbers of lateral-line scales differ slightly. About 66 per cent
of the 149 N. xanthicara counted have 34 scales in the lateral series; 21 per
cent has 33 (mean 33.9, range 33-36). In the sample of 447 rutilus,
about 42 per cent has 34 scales, and 40 per cent has 33 (mean 33.6,
range 32-36). The holotype of xanthicara has 35 lateral-line scales, and
lacks a lateral canal on the three posterior scales on the left side; al-
though some scales are regenerated, the canal appears normal on the

496 Proceedings of the Biological Society of Washington

right side and such aberrations are rare in the series examined. The
lateral line is gently decurved in xanthicara, and there is little pigmenta-
tion associated with it except where it passes through the discrete lateral
band. N. rutilus has a strongly decurved lateral-line canal, and melano-
phores are often positioned as small spots or crescents along the canal at
scale margins.

The head of the new form is attenuate, slender dorso-ventrally, and its
length averages 26.6 per cent of standard length (23.1-29.1; 24.8-28.0).
Head length in rutilus is similar, averaging 26.1 per cent (22.9-30.2; 24.7—
28.8), but it is more blunt (Fig. 1). These slight differences reflect in
part the longer snout of xanthicara, 8.0 per cent of standard length (6.2-
9.5; 6.8-8.8), as opposed to an average of 7.1 per cent for rutilus (5.3-
9.0; 6.6-8.2). Orbital lengths are very similar, 9.0 per cent (6.4-11.9;
8.1-10.6) and 8.4 per cent (6.4—11.3; 7.7-9.3), respectively. The mouths
of both species are oblique, more so in young, and lower jaws are reduced
and included within the upper; the angles from the mandibles to the
lower side of the head are abrupt (Fig. 1).

Body depth and width of N. xanthicara, respectively, average 21.7 per
cent (17.8-25.6; 19.1-23.3) and 12.4 per cent (10.0-14.9; 11.1-13.5) of
standard length, as opposed to 23.1 per cent (18.9-28.1; 21.3-25.5) and
12.2 per cent (8.6—-17.4; 10.5-14.3) for rutilus. Variation is high as a re-
sult of sexual, seasonal, and individual variations. The slenderer body of
xanthicara is reflected, however, in the less variable measurement from
the dorsal origin to the anal origin, which averages 25.3 per cent for the
new form (21.2-27.8; 23.5-26.3) and 27.0 per cent for rutilus (23.2—
32.8; 25.3-29.1). The discrete lateral band of xanthicara emphasizes its
slightly slenderer body and gives the impression of an elongate fish (Fig.
1). Depth of caudal peduncle averages 10.3 per cent (8.9-13.2; 9.6-
11.0) of standard length, and its length averages 23.1 per cent (18.8—
26.1; 21.7-24.0) in xanthicara; rutilus has a thicker, slightly longer caudal
peduncle, depth 11.1 per cent (9.3-15.3; 11.0-12.4) and length 24.1 per
cent (20.3-28.2; 23.2-25.1).

Pigmentation of N. xanthicara is highly diagnostic. The external aspect
is dominated by a discrete, blue-black lateral band that extends from the
darkened two to five (usually four) rays of the middle caudal fin, through
a diffusely-broadened caudal spot, to the back of the eye, becoming varia-
bly diffuse, downward, onto the opercle. The cornea is relatively unpig-
mented, but the band continues as a pre-ocular bar that terminates about
2/3 of the way along the snout. The lateral band of rutilus is much less
discrete, and is highly variable in expression. Slight blackening of
the central caudal fin-rays in rutilus is separated from the lateral band by
a depigmented area just over the end of the hypural plate. The band
usually broadens slightly below the dorsal-fin base, and it may almost dis-
appear as it passes anteriad, especially in breeding individuals. The lateral
pigmentation is scarcely evident above the opercles of rutilus, and the
preocular component usually is obscured by darker snout pigmentation.

New Mexican fish 497

The body of N. xanthicara is pallid except for the lateral band. Scale
margins on the dorsum are diffusely pigmented, giving a cross-hatched
appearance (intensive in breeding individuals), but melanophores are
scarce and randomly distributed on the ventro-lateral surfaces. The
belly is surficially immaculate, with the peritoneum showing through at
the midline. Cross-hatching on the upper sides and back of rutilus is ac-
companied by pigmentation of the intervening spaces, giving a much
darker aspect. Melanophores often extend below the lateral band, espe-
cially onto the abdomen. The belly of rutilus is white, and the usually-
speckled peritoneum does not show through at the midline. The pig-
mentation of the peritoneum of rutilus ranges from dusky to speckled, but
it is invariably black or dark brown in xanthicara.

The predorsal streak of N. xanthicara is broad and diffuse; a postdorsal
streak often is lacking. The predorsal streak is interrupted by a narrow,
relatively depigmented band at the nape, marking the passage of the
supratemporal canal, then broadens to a heart-shaped or sub-hexagonal
spot over the parietal and posterior part of the frontal bones. This spot
is bounded anteriorly by another, transverse, elliptical, depigmented area.
There is a second, heart-shaped area of large melanophores over the fron-
tal bones, between the orbits, then fine melanophores dust the snout and
(variably) the upper part of the upper lip. The arc of the lower part of
the lower lip is similarly dusted with melanin, and this leads medially into
a dusky gular streak. The streak may involve the extreme anterior part of
the breast in some darker fish, but it is sometimes absent in lightened in-
dividuals. Except for a circumorbital scattering of melanophores, and
an occasional larger one, the cheeks, lower parts of the branchiostegal
rays, and the medial parts of the mandibles are silvery white.

Both a predorsal and a postdorsal streak are present in N. rutilus, the
latter less distinct. Details of dorsal head pigmentation are typically
masked by dark melanophores that may extend to the tip of the snout
and to the tip of the upper jaw. There is strong circumorbital melaniza-
tion. The gular streak is typically dark, and extends back to include the
anterior part of the breast in some individuals.

There are deep-lying melanophores associated with the bases of both
the anal and dorsal fins in both species. However, these are often masked
by other pigment in N. rutilus. Pigmentation of the fins themselves in
non-breeding individuals of xanthicara is mostly restricted to melano-
phores lining each delicate ray. The dorsal fin has a dark anterior margin,
resulting from greater concentrations of pigment along the leading rays.
In some fish elongate melanophores darken the interradials of the distal
third of this fin. There is a tendency for concentration of pigment on the
procurrent caudal fin rays of xanthicara, which, along with the caudal
extension of the lateral band, gives an impression of a distinct triad of
caudal spots in individuals with more intense pigmentation. The anal
fin is sparsely pigmented, except at its base. The pelvic fins have a
darkened leading edge, but the remainder is clear. The pectoral fins are
blackened anteriorly, with scattered melanophores posteriorly, and distally

498 Proceedings of the Biological Society of Washington

darkened on rays and interradials in some individuals. Pigmentation of
the fins of rutilus is exaggerated over that of xanthicara, especially in the
dorsal fin which often appears sooty gray. The procurrent and major
unbranched caudal fin-rays are blackened, but no pattern is evident.

Breeding males of N. xanthicara are exceedingly colorful. The follow-
ing notes, transcribed in edited form from field observations of the holo-
type and male paratopotypes, span most variations that we have seen:
head brassy-gold over dorsum and onto snout, color interrupted on sides
of snout and on opercles by dark melanophores, but continuing ventrally
on opercle; yellow continuing on dorsum to caudal fin, overlying gray
or brown ground cclor that is distinctly cross-hatched. Lateral band in-
tensely prominent; sides below lateral band pinkish-orange; belly white,
slightly suffused with yellow; pectoral fins lemon-yellow in center, color
more intense anteriad and with a dark black, leading edge; pelvic fins
with a dark leading edge margined posteriad by an irridescent, bluish
line (milky-white in some), then the remainder of the fin yellow; anal
fin transparent distally and proximally, yellow centrally; caudal fin in-
tense yellow in belly of lobes, light yellow proximally, with procurrent
rays and central rays black; dorsal fin yellow-orange, opaque, with milky-
white pigments proximally, edged with black; cornea yellow, reflecting
blue dorsally; pupil jet black.

Breeding colors of male N. rutilus are to be detailed elsewhere. For
purposes of comparison they consist of a greenish-yellow over-all aspect
on the body, more intensely yellow below. The opercles and sides of the
head are yellow and the dorsum of the head and snout are green or
orange-green (the latter is rare). The fins are milky-yellow to bright
yellow, except for the dorsal which sometimes is blackened. Very little
black pigmentation is evident, except in the dorsal fin, and the lateral
band is often totally masked.

Tuberculation of breeding males of both species consists of strong
organs on the dorsum of the head, separated from smaller ones on the
snout by a narrow, pre-narial hiatus. Tubercles often appear on the chin,
and granulations are present on the cheeks and gular areas. N. rutilus
usually develops small tubercles on the nape, while xanthicara does not.
There is a strong band of densely concentrated tubercles above the anal
base and along the lower sides of the caudal peduncle in both forms. This
rarely extends more than half the length of the peduncle in rutilus, but
may go to the procurrent rays of the caudal fin in xanthicara. Tubercles
develop on all fins, those on the pectorals usually as a double row on the
second through fifth rays and as single rows on the sixth and (rarely)
seventh rays. On other fins, tubercles vary in occurrence on the second
through fourth or fifth rays; they are rarely present on the dorsal or caudal
fins.

Etymology: The name “xanthicara” is a compound of the transliter-
ated Greek words “xanthos” (yellow) and “cara” (or “kara”; top of head),
that describes part of the breeding coloration of the new form. “Kara” is
a neuter noun; its use in a compound name dictates retention of its end-

New Mexican fish 499

ing. We, however, follow general usage (Hubbs and Hubbs, 1958) in
treating Notropis as masculine.

Discussion: N. xanthicara presently is allopatric to other species of
Notropis. Intensive collecting in the Rio Salado de los Nadadores, just
east of the Cuatro Cienegas basin, and elsewhere, has produced only N.
rutilus. Conversely, samples from the canals on the north and northeast
(outlet) sides of the basin have produced no specimens of either species,
and xanthicara prevails (though rare) in parts of the basin where rutilus
might be expected. E. G. Marsh, Jr. collected a Notropis at two places
within the Cuatro Ciénegas basin in 1939, most likely from Canal de La
Angostura (Minckley, 1969). These specimens (UMMZ 130377, 130387 )
correspond with rutilus in all critical characters of pigmentation, and in
features of body proportions and meristics. In more than 50 seining col-
lections in that canal since 1960, Notropis was taken only once, and they
are xanthicara (KU 7374). As noted above, shiners are very rare on the
north side of the basin. Perhaps more intensive manipulation of canals
in that area now than in the past excludes these fishes.

Some specimens of N. rutilus from the Rio Salado de los Nadadores
have pigmentation approaching that of xanthicara. This is especially evi-
dent in non-breeding fish. We have been unable to define intermediacy of
these samples on the basis of other characters, and presently consider this
a response to the clarity of water in that stream. However, it is possible
that hybridization influence, the selective introgression of pigment charac-
ters, has occurred between xanthicara and the downstream population of
rutilus; the problem is under additional study.

Five specimens from Laguna de los Fresnos (Santa Tecla system; KU
7421) differ somewhat from other N. xanthicara. They tend to have long
heads (27.2 per cent of standard length, range 25.4—28.6), with reduced
snouts (7.4 per cent, 6.8-7.9) and large eyes (10.2 per cent, 9.4-11.7).
Their body depths and the measurement from dorsal to anal-fin origin
are high (22.6 per cent [21.1-25.1] and 27.0 per cent [25.5-28.7], re-
spectively ), tending toward rutilus. Pigmentation of these fish is darker
than usual for xanthicara, and is much more similar to that species than to
rutilus; their peritonea are black. The Santa Tecla system is largely dis-
junct at present, partially a result of canalization, and has been studied less
than other parts of the Cuatro Ciénegas basin. Nevertheless, 17 collections
have been made in the system, and only one included shiners. We con-
sider the Los Fresnos specimens to be a form of xanthicara, deferring
speculation on their status until additional material becomes available.

There seems little doubt that N. xanthicara is derived from N. rutilus or
its progenitor, probably in isolation in the large springs and spring-fed
streams of the basin it now inhabits. Isolation of the Cuatro Ciénegas ba-
sin has undoubtedly occurred sporadically for millenia; levels of differen-
tiation of aquatic organisms living there range from endemic subfamilies
and genera in molluscs (Taylor, 1966), through vertebrates and inverte-
brates that do not differ from animals living outside (Minckley, 1969).

500 Proceedings of the Biological Society of Washington

N. xanthicara is intermediate to these extremes, and probably dates to
later Pleistocene. The other five described endemic fishes range from
Cyprinodon bifasciatus Miller (1968), which is considered an ancient,
pre-Pleistocene, relict, through the distinct, but less highly differentiated
C. atrorus Miller, Lucania interioris Hubbs and Miller (1965), Gambusia
longispinis Minckley (1962), and Xiphophorus gordoni Miller and Minck-
ley (1963). Three cichlids and a darter (Percidae) remain to be de-
scribed, which will elevate the total endemic fish component of the basin
to more than 50 per cent (10 of 18 known species). Other species, a cat-
fish, cf. Ictalurus lupus (Girard), a largemouth bass, Micropterus sal-
moides (Lacépéde), and a sunfish, Lepomis megalotis (Ratinesque ), all
are different from the forms outside the basin, though probably not more
than subspecifically.

N. xanthicara occurs widely in the Cuatro Ciénegas basin (Fig. 2), but
has never been taken in shallow marshes or in larger, terminal lakes. It
is most abundant in the upper reaches of streams, just below their
origins in limnocrenes, and often concentrates at the transition dividing
lentic and lotic conditions. In streams and canals the fish lives in groups
of three to 25 individuals in zones of shear between current and back-
water, or moves near the bottom in current. In day time they are rarely
on bottom, but forage at the surface and inspect almost all floating ob-
jects at any depth. At night, the fish rest on the bottom in eddies, in
groups of four to more than 30 individuals, and are lethargic in the spot-
light of a diver. Nothing has been observed on the breeding activities of
this species.

N. rutilus tends to concentrate above and below riffles, with aggrega-
tions of nuptial males often occurring on the swiftest riffles and females
remaining in associated eddies or pools.

The most common associates of N. xanthicara are the Mexican tetra,
Astyanax fasciatus mexicanus (Filippi) and Dionda episcopa Girard, the
roundnose minnow, both in the springs and in streams. Both other species
tend to favor swifter waters, however, and are more active and aggressive.
At one time or another, N. xanthicara has been caught in association with
most other fishes in the basin (Minckley, 1969), except for those that
characterize marshes or saline lakes (C. atrorus, L. interioris, and G.
longispinis ).

LITERATURE CITED
Gipss, R. H., Jr. 1957. Cyprinid fishes of the subgenus Cyprinella of
Notropis. I. Systematic status of the subgenus Cyprinella,
with a key to the species exclusive of the lutrensis-ornatus
complex. Copeia 1957: 185-195.
Husss, C. L. ann C, Husss. 1958. Notropis saladonis, a new cyprinid
fish endemic in the Rio Salado of northeastern México. ibid.
1958: 297-307.
and R. R. Minter. 1965. Studies of cyprinodont fishes.
XXII. Variation in Lucania parva, its establishment in west-
ern United States, and description of a new species from an

New Mexican fish 501

interior basin in Coahuila, México. Misc. Publ. Mus. Zool.,
Univ. Michigan 127: 1-104, 3 pls.

Husss, C. 1954. Corrected distributional records for Texas freshwater
fishes. Texas Jour. Sci. 6: 277-291.

Koster, W. J. 1957. Guide to the Fishes of New Mexico. Univ. New
Mexico Press, vii + 116 pp.

Mitter, R. R. 1968. Two new fishes of the genus Cyprinodon from the
Cuatro Ciénegas basin, Coahuila, México. Occ. Pap. Mus.
Zool., Univ. Michigan 659: 1-15.
AND W. L. Mincxuiey. 1963. Xiphophorus gordoni, a new
species of platyfish from Coahuila, México. Copeia 1963:
538-546.

MiIncKLEY, W. L. 1962. Two new species of fishes of the genus Gam-
busia (Poeciliidae) from northeastern México. ibid. 1962:
391-396.
1969. Environments of the Bolsén of Cuatro Ciénegas, Coa-
huila, Mexico, with special reference to the aquatic biota.
Publ. Univ. Texas E] Paso, Sci. Ser. 2: in press.

Taytor, D. W. 1966. A remarkable snail fauna from Coahuila, México.
Veliger 9: 152-228, 11 pls.

502 Proceedings of the Biological Society of Washington

v=

Vol. & 17 November 1969

PROCEEDINGS
OF THE

BIOLOGICAL INVESTIGATIONS OF THE DEEP SEA.
50. THE VALIDITY AND GENERIC POSITION OF
PENTAGONASTER PARVUS PERRIER (ECHINODER-
MATA, ASTEROIDEA )!

By JeraLtp A. HALPERN
Institute of Marine Sciences, University of Miami

When Perrier reported on the sea stars collected by the
Blake (1881, 1884), one of the new species he described was
Pentagonaster parvus. Verrill (1899: 151-156) examined the
syntypes of Pentagonaster parvus and concluded that they were
young specimens of Goniaster americanus Verrill (= Asterias
tessellatus Lamarck). I have examined the syntypes of P.
parvus, as well as many other specimens. I have also examined
many young specimens of Goniaster tessellatus and have con-
cluded that Pentagonaster parvus is a valid species belonging
to the genus Tosia.

This research was supported by the National Science Founda-
tion through grant GB-4936. I am grateful for the assistance of
Dr. Lowell P. Thomas of the Institute of Marine Sciences,
University of Miami, Dr. H. B. Fell of the Museum of Com-
parative Zoology, Harvard and Miss Maureen E. Downey of
the U.S. National Museum.

Tosia parva (Perrier, 1881)

Pentagonaster parvus Perrier, 1881, p. 19; 1884, pp. 36, 37, 231, pl. 7,
figs. 7-8.—Sladen, 1889, pp. 265, 267, 746-747.—?H. L. Clark,
1898, p. 5.

Goniaster americanus (pars) Verrill, 1899, pp. 154-156, pl. 26, fig. 6.

Plinthaster dentatus (pars) Gray, et al., 1968, fig. 25.

Material studied: 22 specimens from the following localities: Lecto-
type: R = 10.5 mm, r = 7.5 mm, R/r = 1.4; Blake sta. 253, off Grenada,

1 Contribution No. 1098 from the Institute of Marine Sciences, University of Miami.

41—Proc. Bion. Soc. Wasu., Vou. 82, 1969 (503 )

504 Proceedings of the Biological Society of Washington

Fic. 1. Tosia parva (Perrier). Top, specimen from Pillsbury sta. 707,
abactinal view, 6.1.—Bottom, specimen from Silver Bay sta. 2263, ac-
tinal view, 7.8.

Pentagonaster parvus 505

168 m, 1878-79, MCZ 417.—Paralectotypes: Blake, West Indies, 172—
229 m, 1877-79, MC 4283, (3 spec. ).—1 spec., M/V Silver Bay sta. 2263,
33°04’N, 78°12'W, 30 m, 28 July 1960, UMML 40.149.—1 spec., R/V
Hernando Cortez sta. E, 27°36'N, 84°13’W, 73 m, 4 January 1966,
USNM E10851.—1 spec., M/V Silver Bay sta. 3496, 20°53’N, 73°42’W,
183, 4 November 1961, UMML 40.159.—1 spec., R/V Pillsbury sta. 478,
11°33'N, 62°09'W, 597 m, 2 August 1966, UMML 40.235.—2 spec., R/V
Pillsbury sta. 707, 11°22’N, 62°22’W, 79 m, 19 July 1968, UMML
40.236.—13 spec., R/V Pillsbury sta. 734, 11°01'N, 63°35’W, 60-71 m,
22 July 1968.

Diagnosis: R not greater than 30 mm; R/r = 1.3-1.8. Abactinal and
marginal plates bearing scattered granules in centers. Peripheral granules
of abactinals in radial areas fused. Actinals surrounded by more than
single row of granules. Five compressed adambulacral furrow spines;
nine to ten mouth furrow spines.

Description: Five arms. R = 22 mm, r = 46 mm, R/r = 1.5. General
from pentagonal to arcuate pentagonal.

Five primary plates conspicuously larger than other abactinals. Abac-
tinal plates slightly convex; each surrounded by single row of large,
flattened, rectangular granules. Granules surrounding plates in radial
areas fused so that each plate surrounded by flattened, calcareous ring.
Center of each abactinal plate bearing one to six round granules em-
bedded in deep pits. Papulae confined to large radial areas.

Inferomarginals and superomarginals corresponding throughout wide
interbrachial arc. Eight massive superomarginal plates; each surrounded
by single row of small, rounded granules. Clusters of one to twelve round
granules, similar to those of abactinals, scattered about each plate. Double
row of large, flattened granules, similar to those surrounding abactinals
along suture between superomarginals and inferomarginals. Granules
twice as large at angle formed by two adjacent superomarginals and cor-
responding inferomarginals. Each enlarged granule bearing single, very
small, rounded granule. Terminal plate small, naked. Ten massive in-
feromarginal plates; granulation similar to superomarginals.

Actinal intermediate area large; plates arranged in five chevrons.
Actinal plates large, flat, rhombic; each plate surrounded by two to three
irregular rows of coarse, rounded granules. Most plates having large naked
central area; some plates bearing one to four scattered granules in central
area.

Adambulacral plates square with straight furrow margin bearing five
subequal, compressed furrow spines with blunt, rounded tips. Sub-
ambulacral spines in three to four irregular rows of three to five short,
blunt spinules slightly taller than granules of actinal plates.

Each mouth plate bearing nine to ten furrow spines, similar to adam-
bulacral furrow spines, but more strongly compressed, median spine being
most compressed; median spine only slightly enlarged. Rest of plate
covered by ten to twelve pyramidal spinules, slightly taller than subam-
bulacral spines of adambulacrals.

506 Proceedings of the Biological Society of Washington

Anus prominent, subcentral. Madreporite roundly triangular, about
two-thirds as large as adjacent abactinal plates; located approximately one-
third the distance from center of disk to middle of interbrachial arc. No
pedicellariae.

Type: Museum of Comparative Zoology, cat. no. 417 (lectotype ).

Type-locality: off Grenada, 168 m, Blake sta. 253.

Distribution: This species is found throughout the Antillean province,
extending from 50 miles south of Cape Fear, North Carolina to Trinidad.
Its bathymetric range is 30-597 m.

Discussion: The smallest specimen examined measures R = 6 mm. All
its characters are the same as in an adult, except the peripheral granules
of the abactinals of the radial areas are not yet tused.

The smallest specimen of Goniaster tessellatus I examined (R = 9 mm)
is distinguished from Tosia parva by having naked superomarginals and
the abactinal and actinal plates completely covered by granules. Specimens
as small as R = 11 mm already have abactinal spines forming.

Pentagonaster parvus Perrier belongs in Tosia because of its pentag-
onal form, actinal granulation and lack of pedicellariae. It is distin-
guished from all other species of Tosia by its central abactinal and mar-
ginal granules. It is the only species of Tosia not from Australian waters
and is a Tethyan relict.

The specimen collected at Blake station 253 is designated the lecto-
type and the type locality is restricted to off Grenada, 168 m. The speci-
mens collected by the Blake at stations 32, 276 and 296 have been placed
together and it is impossible to determine which specimen is from which
station. These three specimens are designated the paralectotypes.

LITERATURE CITED

Crark, H. L. 1898. The echinoids and asteroids of Jamaica. Johns
Hopkins Univ. Cire., 18 (137): 4-6.

Gray, I. E., M. E. Downey, AND M. J. CerRAME-VivAs. 1968. Sea-stars
of North Carolina. Fish. Bull., 67 (1): 127-163, figs. 1-40.

Perrier, E. 1881. Reports on the results dredging . . . in the Gulf of
Mexico, and in the Caribbean Sea. 1877-79, by the U.S.
Coastal Survey steamer “Blake”. . . 14. Description sommaire
des espéces nouvelles d’Astéries. Bull. Mus. comp. Zool.
Harvard, 9 (1): 1-31.
1884. Mémoire sur les étoiles de mer recueillies dans la Mer
des Antilles et la Golfe du Mexique durant les expéditions de
dragage faites sous la direction de M. Alexandre Agassiz.
Nouv. Arch. Mus. Hist. nat. Paris, (2) 6: 127-276, pls. 1-9.

SLADEN, W. P. 1889. Report on the Asteroidea collected by H. M. S.
CHALLENGER during the years 1873-1876. Reports Chal-
lenger Zool., 30: 1-893, pls. 1-117.

VerritL, A. E. 1899. Revision of certain genera and species of star-
fishes. Trans. Conn. Acad. Arts, Sci., 10: 145-234, pls. 24—
30.

4 7

ww

Vol. 85 ak 17 November 1969

PROCEEDINGS
OF THE

pIULULILAL SOCIETY OF WASHINGTON

A NEW SPECIES OF CAPRELLID (CRUSTACEA:
AMPHIPODA ) FROM OREGON

By Joun C. McCain
Smithsonian Oceanographic Sorting Center, Washington, D.C.

In my paper on the Caprellidae of the Western North At-
lantic (1968, p. 52) I discussed the species then known to be
associated with echinoderms. Four species were listed which
had been found clinging to sea stars. A fifth, belonging to an
undescribed species, was brought to my attention by Irwin
Polls and Dennis S. Greenley of Oregon State University. This
species is herein described as new and named in honor of
one of the collectors of the holotype.

Caprella greenleyi new species
Figure 1

Material examined: Oregon, Boiler Bay, collected by Irwin Polls and
Dennis S, Greenley, intertidal from the starfish Henricia leviuscula, 1 ¢
holotype USNM 123523, 1 ovig. 2 allotype USNM 123524, 1 9 para-
type USNM 123525.

Description: Male holotype: Body and appendages robust and covered
with microtubercles, pereonites 3 and 4 with anteriorly directed pleural
projections. Length 2.7 mm.

Antenna 1 approximately length of pereonites 1 and 2, flagellum uni-
articulate. Antenna 2 slightly shorter than antenna 1, flagellum uni-
articulate.

Mouthparts typical of Caprella (McCain, 1968, p. 18).

Gnathopod 1 typical of genus, grasping margin of dactylus and propo-
dus serrate, propodus with 2 proximal grasping spines. Propodus of
gnathopod 2 broad, approximately 2/3 as wide as long, grasping margin
with 2 proximal grasping spines and medial notch; dactylus massive and
scimitar-shaped.

Propodus of pereopods 5—7 with 2 proximal grasping spines.

Abdomen typical of genus.

42—Proc. Biot. Soc. WAsH., Vou. 82, 1969 (507)

508 Proceedings of the Biological Society of Washington

Ficure 1. Caprella greenleyi new species. Male holotype: a, lateral
view; b, gnathopod 1; c, antenna 2; d, gnathopod 2; e, pereopod 7.

Remarks:

This species can be distinguished from the other species of Caprella
due to its small size, 2.7 to 3.6 mm, and the large broad propodus of
gnathopod 2 with a massive, scimitar-shaped dactylus.

The male holotype and an ovigerous female of 3.6 mm were found
clinging to a specimen of Henricia leviuscula which measured 6 cm from
disk to arm tip. The female paratype measured 3.5 mm and was col-
lected from a smaller Henricia of 1.6 cm. The caprellids were noticed

New Oregon caprellid 509

after the sea stars were returned to the Yaquina Marine Biological Lab-
oratory where they were held in tanks for study. A field examination of
numerous specimens of Henricia failed to yield any other specimens of
C. greenleyi.

LITERATURE CITED

McCain, Joun C. 1968. The Caprellidae (Crustacea: Amphipoda) of
the western North Atlantic. Bull. United States Nat. Mus.,
vol. 278, pp. vi + 147, 56 figs.

510 Proceedings of the Biological Society of Washington

Vol. 17 November 1969
PROCEEDINGS
OF THE

BIOLOGICAL SOCIETY OF WASHINGTON

SYRINGONOMUS TYPICUS NEW GENUS, NEW SPECIES
(ENOPLIDA: LEPTOSOMATIDAE) A MARINE
NEMATODE INHABITING ARENACEOUS TUBES.

By W. Duane Hope anp D. G. Murpuy*
Smithsonian Institution, Washington, D.C.

Several collections of marine nematodes have been obtained
from epibenthic trawls taken by the Woods Hole Oceano-
graphic vessel, ATLANTIS II, on transects from Woods Hole,
Massachusetts to Bermuda. Numerous specimens in one of
these collections were partially enclosed in tubes, the latter
usually cylindrical in shape and constructed of adhering par-
ticles of sand. The lengths of the tubes vary considerably due
in part to breakage, but each has similar construction, and,
with few exceptions, each accommodated a single nematode.
Thirty-nine nematodes were removed from tubes and examined
more carefully. Of this number, six are of species as yet uniden-
tified. The remaining 33 are males, females and juveniles of a
new genus and new species described below:

Syringonomus new genus

Diagnosis: Same as that of Leptosomatinae Filipjev, 1916. Body
slightly tapered anteriorly and posteriorly. Cuticle smooth. Tail bluntly
conical. Cephalic setae short. Cephalic capsule present, but apparent
only in optical section. Amphid a small, indistinct pore. Stoma un-
armed. Eyespots absent. Gubernaculum consisting of small, paired struc-
tures, lateral to distal ends of spicula; corpus of gubernaculum and lateral
anterior projection absent. Setiform subventral supplements present, ven-
tromedian supplements absent. Caudal glands and spinneret present.

Etymology: The name Syringonomus is derived from the Greek Syrin-
gos meaning tube, and nomos meaning a place for living.

*Visiting Research Associate.

43—Proc. Brot. Soc. WaAsu., Vou. 82, 1969 (511)

512 Proceedings of the Biological Society of Washington

Syringonomus typicus new species

Specimens: Holotype (Male): National Museum of Natural History Num-
ber 39489.
Allotype (Female): National Museum of Natural History Number
39493.
Paratypes (Males): National Museum of Natural History Numbers
39490 thru 39492.
Paratypes (Females): National Museum of Natural History Num-
bers 39494 thru 39515.
Paratypes (Juveniles): National Museum of Natural History Num-
bers 39516 thru 39539.
Measurements:
Holotype: L = 5.377 mm; a = 65.9; b = 7.3; ¢ = 37.9
Allotype? lL: = 4,936 mm: @ = 40.1; b = 7.1; ¢ = 41-8; V =] Glo.
Male Paratypes*: L = 6.061 mm; a = 64.1; b = 8.5; c = 42.7.
L = 5.979 mm; a = 64.6; b = 8.3; c = 48.6.
Female Paratypes:

L= 3.32 -4.94 mm (4.23 mm + 0.51 mm)
a= 31.5 - 46.4 (38.5 + 4.3)
b=] 54= 71 (6.3 + 0.6)
c= 23.1-51.0 (40.8 + 7.9)
V — 51.6 - 66.2 (63.4 + 3.8)

Description: Body slender and gradually tapering anteriorly (Fig.
1B); posteriorly, body of nearly uniform diameter to level of anus, then
tapering to form bluntly conical tail (Figs. 2A and B). Head diameter at
level of cephalic setae 30.0 w — 34.7 w (32.3 4 + 1.5 w). Body diameter
at base of esophagus 71.0 » — 89.5 w (82.6 w + 7.3 w) in males, 76.5 uw -
102.0 uw (91.6 « + 7.4 u) in females**; at mid-body length 81.5 u—96.0 u
(91.6 uw + 5.8 uw) in males, 83.0 » — 126.0 » (110.5 » + 13.7 p») in fe-
males; at level of anus 76.5 « — 100.0 u + 6.7 uw).

Cuticle smooth. Head with circle of six cephalic papillae and second
circle of 10 cephalic setae (Figs. 1A and B); longer cephalic setae 4.0 » —
6.0 uw, shorter 2.5 « — 4.7 uw. Distance from anterior extremity of head to
level of cephalic setae 12.0 uw — 19.0 uw (14.3 w + 2.1 w). Somatic setae
equally short and sparse. Amphid an obscure circular pore approximately
1.0 uw in diameter, located 14.4 w — 23.3 uw (19.6 » + 2.3 «) from anterior
extremity of head (Figs. 1A and C). Males with inverted lyre-shaped
pattern on cuticle immediately posterior to amphid; pattern crenate and
with (Fig. 1C) or without posteriorly directed central process. Cuticle
thickened at level of pattern (Fig. 1C). Cephalic capsule present, but
situated anterior to cephalic setae and visible in optical section only
(Figs. 1A, B, and C).

* One male sectioned.

** Measurements of males are given separate from those of females only where the
mean values appear to differ significantly; otherwise measurements of males and
females are combined.

A new marine nematode 513

Fic. 1. Syringonomus typicus new species. A. Lateral view of female
head (allotype). B. Lateral view of head and neck of female (allotype).
Ventral gland cell, VGC. C. Lateral view of male head (holotype).

514 Proceedings of the Biological Society of Washington

ri ae ae

Fic. 2. Syringonomus typicus new species. A. Lateral view of female tail
(allotype). B. Subventral view of male tail (holotype). Spiculum, SP;
Gubernaculum, GU. C. Anterior end of specimen extending from arena-
ceous tube.

A new marine nematode ol5

Head rounded without lips or microlabia. Stoma narrow, not morpho-
logically distinct from lumen of esophagus. Teeth absent (Figs. 1A and
C).

Some specimens with indistinct duct and pore of ventral gland (Fig.
1B), apparently absent in others. Distance from anterior extremity of
head to ventral gland pore 15.0 — 24.6 (19.2 + 4.0) per cent of esopha-
gus length.

Esophagus cylindrical, 628 u — 790 u (717 » + 58 w) long in males,
556 wu — 717 w (664 uw» + 45 w) in females. Eyespots absent. Pseudocoelom
with large, lobate cell on each lateral side of esophagus base.

Caudal glands outstretched and extending anterior to rectum. Cuticle
of tail terminus with median, crescent-shaped lamella. Caudal gland pore
slightly ventral to terminus. Caudal setae sparse, terminal setae absent.

Males—Diorchic, testes opposed and outstretched. Spicula paired,
equal in length, slightly arched, and 72 uw to 75 uw long. Gubernacula
small, tube-like structures, one lateral to distal end of each spiculum;
apophyses and lateral anterior projections absent (Fig. 2B). Dorsoven-
tral copulatory muscles sparse, posterior region of body not curved ven-
trally. Each side of body with two to four setiform subventral supple-
ments; setae approximately 3 uw long, first pair 30 uw to 37 uw, second 113 pu
to 120 u, third 132 uw and fourth 149 w anterior to cloacal vent; setae fur-
thest anterior slightly closer to ventromedian line. Ventromedian supple-
ments absent. Tail length 120 1 — 142 u (13lu+104.z).

Females—Didelphic, gonads opposed and reflexed; vulva 1.87 mm —
3.17 mm (2.60 mm-—0.41 mm) from anterior end. Tail length 85 u—-
1444 (106 u+ 164).

Type Locality: Sediment from epibenthic traw] taken between 39°
37.0’ N, 66° 47.0’ W and 39° 37.5’ N, 66° 44.0’ W at 3,806 meters depth
on 24 August, 1966.

Discussion: Specimens of Syringonomus typicus possess characters typi-
cal of the subfamily Leptosomatinae. They most closely resemble species
of the genera Leptosomella Filipjev, 1925, Leptosomatides Filipjev, 1918,
Paraleptosomatides Mawson, 1956, Leptosomatina Allgen, 1951, and
Leptosomatum Bastin, 1865. Leptosomella differs in having long cephalic
setae and an acutely conical tail. Leptosomatides and Paraleptosomatides
differ in having complex gubernacula, supplements, and well developed,
setiform, subventral supplements, the more anterior ones on cuticular
elevations. Leptosomatina differs in having long cephalic setae, armed
stoma, and complex gubernaculum with caudally directed apophyses.
Finally, Leptosomatum, whose members most closely resemble Syringono-
mus, differs in not having the lyre-shaped pattern and thickened cuticle
on the head of the males. By the latter two characters, Syringonomus may
be distinguished also from all other genera of this subfamily.

The presence of a ventral excretory cell is insufficiently documented to
be relied upon at this time as a diagnostic character.

A striking feature of the specimens under consideration is that they
were found inhabiting hollow, cylindrical tubes constructed of sand parti-

516 Proceedings of the Biological Society of Washington

cles and an adhesive mortar. The lengths of the tubes range from 1.0 mm
to 3.0 mm, and the width from 0.5 mm to 2.0 mm. The diameter of the
sand particles in the tubes range from 138 uw to 588 uw with an average
diameter of 362 u. The average diameter of sand particles from the tube
constructed of the finest sand was 189 yu, and 428 wu in the case of the
tube constructed of the coarsest particles. The particles are primarily
quartz.

The lumen of each tube is lined with a thin layer of what is presumed
to be identical to the mortar between sand particles. The lining varies from
light yellow in some tubes to dark brown in others. The lining and mortar
become dark blue when treated with equal volumes of 2. percent hydro-
chloric acid and 2 percent postassium ferrocyanide demonstrating both
contain ferric compounds.

Of particular interest is the question of whether or not Syringonomus
typicus is responsible for the construction of the tubes. Obviously, the
organism involved must possess a means of producing the lining and mor-
tar. Many marine nematodes possess caudal glands that secrete an adhesive,
usually employed for attachment to a substrate, and many possess lateral
hypodermal glands, the function of which is as yet unknown. While no
nematodes are known to construct tubes, either or both kinds of glands
could conceivably secrete a substance that would serve as mortar in form-
ing arenaceous tubes. If this were the case, one might expect the glands
involved to be particularly well-developed and perhaps modified in other
respects. However, specimens of Syringonomus typicus do not have what
could be readily identified as lateral hypodermal glands, and while they
do possess caudal glands and a spinneret, they are not exceptionally well-
developed or unusual in other respects. Therefore, while it appears that
this species of nematode is an inhabitant of these tubes, there is little
evidence to suggest they construct them.

Our further attempts to learn the identity of the organism responsible
for construction of the tubes resulted in their being examined by a taxon-
omist of foraminiferans, who identified them as tubes most likely con-
structed by Rhabdammina abyssorum M. Sars, 1868. Descriptions of the
general features of the test of this species are given by Carpenter (1875)
who has found that the test is typically triradiate, the rays diverging at
equal angles from a central cavity and each ray with an orifice at its
extremity. He states further, however, that quadri- and pentaradiate
forms occur as well as single, straight tubes. The latter form “often ex-
ceeds half an inch” in length.

The walls of the test of this species, according to Brady (1884), are
composed chiefly of coarse sand, the grains of which are variable in size.
Brady also found that the walls of tests from the North Atlantic are
various shades of light reddish-brown, and chemical analysis of the mor-
dant demonstrated the presence of peroxide of iron.

The descriptions of the test of this foram closely conform to that of the
tubes inhabited by the nematodes, except that the latter are shorter and
always in the form of a straight tube. It is concluded, therefore, that the

A new marine nematode Bila

specimens of Syringonomus typicus in our collections are within broken
pieces of the tests of Rhabdammina abyssorum. To what extent these
nematodes dwell in these tubes, and to what extent, if at all, they are
ecologically adapted to a tube-dwelling existence, must await further study.

Acknowledgments: The authors wish to express their appreciation to
Mrs. Carolyn Bartlett Gast for preparing the illustrations; to Ruth Todd
of the Geological Survey for identifying the foraminiferan tubes; and to
Dr. Jack Pierce, Department of Paleobiology, Smithsonian Institution, for
determining the diameters of the sand particles.

LITERATURE CITED

ALLGEN, C. A. 1951. Papers from Dr. Th. Mortensons Pacific Expedition,
1914-1916. LXXVI. Pacific Freeliving Marine Nematodes.
Videnskabelige Meddelelser Dansk Naturhistorisk Forening,
Copenhagen 113: 262-411.

Bastian, H. C. 1865. Monograph on the Anguillulidae or Free Nema-
toids, Marine, Land and Freshwater with Descriptions of 100
New Species. Transactions of the Linnean Society of London
25(2): 73-184.

Brapy, H. B. 1884. Report on the Foraminifera collected by H. M. S.
Challenger during the year 1873-1876. Report on the Scien-
tific Results of the Voyage of H. M. S. Challenger during the
years 1873-1876 IX: 1-814. Plates 1-115.

CARPENTER, W. B. 1875. The Microscope and its Revelations. Fifth
Edition. J. and A. Churchill, London. 848 pp.

Finipyev, I. N. 1916. Les Nematodes libres contenus dans les Collec-
tions du Musee Zoologique de Acad. Imp. des Sciences de
petrograd. Extrait de Annuaire du Musee Zoologique de I’-
Acad. Imp. des Sci. 21: 59-116.

1918. Svobodnozhivushchiya Morskiya Nematody Okrest-
nostei Sevastopolya. Trudy osoboi Zoologicheskoi Laboratorii
I Sevastopol ’Skoi Biologicheskoi St. Antsii Rossiiskoi Akade-
mii Nauk, Series II(4): 1-350. (Translated from Russian.
Israel Program for Scientific Translation Jerusalem, 1968).
1925. Les Nematodes libres des mer septentrionales 4 la
famille des Enoplidae. Archiv fiir Naturgeschichte 91: 1-
216.

Mawson, P. M. 1956. Freeliving Nematodes. Section I: Enopoloidea
from Antarctic Stations. British-Australian-New Zealand
Antarctic Research Expedition 6(3): 39-74.

518 Proceedings of the Biological Society of Washington

LV ia
4
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Vol. Ril 17 November 1969

PROCEEDINGS
OF THE

BIOLOGICAL SOCIETY OF WASHINGTON

AUSTRALAUGENERIA POTTSI, NEW NAME FOR
POLYNOE LONGICIRRUS POTTS, FROM THE MALDIVE
ISLANDS (POLYCHAETA: POLYNOIDAE)

By Marian H. PETTIBONE
Smithsonian Institution, Washington, D.C.

The original description of the polynoid polychaete Polynoe
longicirrus Potts, 1910, was based on material collected by Mr.
J. Stanley Gardiner in 1899 from four localities in the Mal-
dive Islands: South Male, North Male (“ off a Gorgonian’),
South Nilandu, and Fadifolu. Syntypes from one of these lo-
calities, that of South Nilandu, are now deposited in the British
Museum (Natural History), having been transferred from the
Cambridge Museum. As pointed out by Augener (1922, p. 10,
footnote) and Hartman (1959, pp. 103, 108, Catalogue),
Polynoe longicirrus Potts, 1910, is a junior homonym of Polynoe
(Lepidonotus) longicirra Schmarda, 1861. In my recent paper
on “A review of some species referred to Scalisetosus McIntosh”
(Pettibone, 1969, p. 25), I indicated that Potts’ Polynoe longicir-
rus might prove to belong to Australaugeneria Pettibone and
that the type-specimens needed to be re-examined. Such re-
examination has now confirmed my earlier supposition and
Potts’ species is herein given a new name and re-described.

I wish to thank David George of the British Museum ( Nat-
ural History ) (BMNH) for the loan of the type-specimens and
Fenner A. Chace, Jr., of the Smithsonian Institution for criti-
cally reading the manuscript. This study was aided in part by
a grant from the National Science Foundation (NSF GB-1269).

FAMILY POLYNOIDAE MALMGREN
Genus Australaugeneria Pettibone, 1969; emended

Type-species: Polynoe rutilans Grube, 1878, by original designation.
Emended diagnosis: Buccal segment (II) without notosetae (type-

44—Proc. Bion. Soc. WasuH., Vou. 82, 1969 (519)

520 Proceedings of the Biological Society of Washington

|
f g hil |

Ficure 1. Australaugeneria pottsi n. name (Syntypes of Polynoe longi-
cirrus Potts, BMNH 1924: 3: 77): a, Dorsal view anterior end, tip of
antenna broken, upper tentacular cirri and first pair elytra missing; b,
elytrigerous parapodium from segment II, anterior view; c, neurosetae
from same; d, cirrigerous parapodium from segment III, posterior view;
e, elytrigerous parapodium from segment IV, anterior view; f, notoseta
from same and tip magnified; g, upper neuroseta from same; h, middle
and lower neurosetae from same.

New name for Polynoe longicirrus 521

species) or notosetae few in number. Presetal neuropodial lobes of seg-
ments II and III enlarged, hoodlike (type-species) or only slightly en-
larged.
Australaugeneria pottsi new name
Figs. 1-3

Polynoe longicirrus Potts, 1910, p. 336, pl. 18, fig. 9, pl. 20, fig. 29, pl. 21,
figs. 37, 38.—Augener, 1922, p. 10: (footnote). Not Polynoe (Lepi-
donotus) longicirra Schmarda, 1861.

Scalisetosus longicirrus (Potts).—Hartman, 1959, p. 108. [HOMONYM.]

Material examined: South Nilandu, Maldive Islands, Indian ocean,
J. S. Gardiner collection—3 syntypes of Polynoe longicirrus Potts (BMNH
1924: 3: 77). [Three anterior fragments of 12, 16 and 20 segments; pos-
terior fragment of 9 segments; and 6 middle fragments. ]

Description: Body small, flattened, tapered gradually posteriorly.
Length 6.5—7.5 mm, width, including setae, 2 mm, segments 37-38. Ely-
tra 15 pairs, arranged on segments 2, 4, 5, 7, alternate segments to 23, 26,
29, and 32. Elytra large, covering dorsum, soft, translucent, smooth,
without tubercles or papillae. Prostomium bilobed, with lobes rounded
anteriorly, without distinct cephalic peaks; ceratophore of median antenna
in anterior notch, with style long and tapered; lateral antenna with dis-
tinct ceratophores, inserted ventrally, with styles very short; ventral palps
short, stout, tapered; no eyes visible (fig. la). Tentacular parapodia (I)
achaetous, with 2 pairs long tentacular cirri. Buccal segment (II) with
ventral buccal cirri slightly longer than following ventral cirri; without
nuchal fold; notopodia small, each with 2 short notosetae; neurosetae
hooked; presetal neuropodial lobe longer than postsetal lobe but not es-
pecially enlarged (fig. la-c). Neurosetae of segments 3 and 4 also more
strongly hooked than following neurosetae (fig. ld-h). Parapodia sub-
biramous (figs. 2a, b, 3a, b). Notopodia small, conical, confined to
middle of neuropodial lobe; notosetae few in number (2-7), short, more
slender than stouter type of neurosetae, slightly curved, with serrated
border and blunt, slightly bidentate tips (figs. 1f, 2c, 3c). Neuropodia
elongate, diagonally truncate distally, deeply notched dorsally and ven-
trally, forming anterior and posterior rounded lobes, former slightly
longer than latter. Neurosetae of 2 types: upper few (2-5), slender,
bent, spinous, with tips blunt (figs. 1g, 2d, 3d); middle and lower neuro-
setae slightly more numerous (6-8), stout, wider subdistally, smooth or
faintly spinous on enlarged part, with slightly hooked tips (figs. 2e,
3e). Dorsal cirri with elongate cylindrical cirrophores and long filamen-
tous styles (figs. 1d, 2a, 3a). Dorsal tubercles inconspicuous. Ventral cirri
short, subulate, extending slightly beyond neuropodial lobes. Two dorsal
transverse ciliated bands per segment.

Distribution: Indian Ocean (Maldives). May be found on gorgonians
(Potts, 1910).

Remarks: A. pottsi differs from the two previously described species of
Australaugeneria from the Philippine Islands and southwest Australia, A.

522 Proceedings of the Biological Society of Washington

st
Ss —
a
BO
a
Ain, Sie L—_—= es Z
ee ee ee
SS —— S

0.1 mm

d e

Ficure 2. Australaugeneria pottsi n. name (Syntypes of Polynoe longi-
cirrus Potts, BMNH 1924: 3: 77): a, Middle cirrigerous parapodium,
posterior view; b, middle elytrigerous parapodium, anterior view; c,
notosetae from same and tip magnified; d, upper neuroseta from same; e,
middle and lower neurosetae from same.

New name for Polynoe longicirrus 523

b

FicureE 3. Australaugeneria pottsi n. name (Syntypes of Polynoe longi-
cirrus Potts, BMNH 1924: 3: 77): a, Posterior cirrigerous parapodium,
posterior view; b, posterior elytrigerous parapodium, anterior view; c,
notoseta from same and tip magnified; d, upper neurosetae from same
and tip magnified; e, middle and lower neurosetae from same.

rutilans (Grube, 1878) and A. michaelseni Pettibone, 1969, in that the
parapodia of segments II and III are less modified, i.e., the presetal
neuropodial lobes are not especially enlarged or hoodlike, the neurosetae
are not as strongly hooked, and two notosetae are present in segment II
and not absent, as in the other two species. A. pottsi agrees more closely
with A. rutilans in having the notosetae more slender than the stoutest
neurosetae, curved, with spinous rows and bifid tips, and not smooth,
stout, spikelike, as in A. michaelseni. The notopodia are short and con-
fined to the middle of the neuropodial lobes in A. pottsi and A. rutilans
and not extending to near the distal tips of the neuropodia, as in A.
michaelseni.

In her Catalogue of the Polychaeta of the World, Hartman (1959, p.
108) referred Polynoe longicirrus Potts, 1910, to Scalisetosus, perhaps
following a suggestion by Augener (1922, p. 10, footnote) that it might
be a Scalisetosus-like form. As indicated by Pettibone (1969), it does
not agree with Scalisetosus McIntosh.

524 Proceedings of the Biological Society of Washington

LITERATURE CITED

AucENER, H. 1922. Revision der australischen Polychaeten-Typen von
Kinberg. Ark. Zool. Stockholm, 14(8): 1-42, 10 figs.

GrusE, E. 1878. Annulata Semperiana. Mém. Acad. Imp. Sci. St. Péters-
bourg, (7), 25 (8): 1-300, 15 pls.

HartTMan, O. 1959. Catalogue of the Polychaetous annelids of the
World. Allan Hancock Found. Publ. Occas. Paper, No. 23:
1-628.

PetTTIBoNE, M. H. 1969. Review of some species referred to Scalisetosus
McIntosh (Polychaeta, Polynoidae). Proc. Biol. Soc. Wash-
ington, 82: 1-30, 12 figs.

Ports, F. A. 1910. Polychaeta of the Indian Ocean. Pt. 2. The Palmy-
ridae, Aphroditidae, Polynoidae, Acoetidae and Sigalionidae.
Trans. Linn. Soc. Zool., ser. 2, 16: 325-353, pls. 18-21.

ScumarpA, L. K. 1861. Neue wirbellose Thiere beobachtet und gesam-
melt auf einer Reise um dei Erde 1853 bis 1857. Lepizig, vol.

1. Turbellarien, Rotatorien und Anneliden. Pt. 2, 1-164, 22
plates, 100 figs.

”

cS

a

ue
NI
i)

Vol. al a 17 November 1969

PROCEEDINGS
OF THE

BIOLOGICAL SOCIETY OF WASHINGTON |

THE POSTLARVAE AND JUVENILE STAGES OF TWO
SPECIES OF PSEUDOSQUILLOPSIS (CRUSTACEA,
STOMATOPODA) FROM THE EASTERN PACIFIC
REGION

By RayMonp B. MANNING
Smithsonian Institution, Washington, D.C.

This report was prompted by a study of the West African
stomatopods in which two gonodactylid postlarvae, 30 to 33
mm in length, were encountered in collections from the Gulf
of Guinea. These specimens resembled members of the genera
Parasquilla and Pseudosquillopsis. One of these had been iden-
tified by Schmitt (1926) as the postlarva (first littoral
stage) of Parasquilla ferussaci { Roux, 1830) although it differs
in several respects from an earlier account of the postlarva of
that species by Giesbrecht (1910).

As a result of comparing these specimens with the postlarvae
and adults of other members of the family, their identity has
been established, diagnostic characteristics of the postlarvae of
Pseudosquillopsis can be summarized, and four species can be
recognized in the genus.

Prior to this study, the genus Pseudosquillopsis was con-
sidered to comprise three species: P. cerisii (Roux, 1828) from
the Mediterranean Sea, P. dofleini (Balss, 1910) from Japan,
and P. lessonii (Guérin, 1830) from the eastern Pacific region
(Seréne, 1962; Manning, 1963). Two additional species, P.
monoceros (H. Milne-Edwards, 1837), from Chile, and P.
marmorata (Lockington, 1877), from California, have been
described, but most authors including Miers (1880), Bigelow
(1894) and Schmitt (1940), synonymized these species with P.
lessonii, usually without comment. However, Miers did note
(1880, p. 114) that “P. marmorata, Lockington (P. Cal. Ac. Sci.

45—Proc. Biot. Soc. WasuH., Vou. 82, 1969 (525)

526 Proceedings of the Biological Society of Washington

cL.

a
a

Ficure 1. Pseudosquillopsis marmorata (Lockington), female post-
larva, TL 26.5 mm: 4a, anterior portion of body; b, raptorial claw; c, lat-
eral processes of sixth and seventh thoracic somites; d, sixth abdominal
somite, telson, and uropod; e, submedian denticles of telson; f, basal pro-
longation of uropod. (Setae omitted ).

p. 33, 1877), from San Diego, California, either belongs to this
species [lessonii] or is very closely allied to it.”

The postlarvae and juveniles treated here clearly show that
two species of Pseudosquillopsis occur in the eastern Pacific
region, and P. marmorata (Lockington ) is recognized as a dis-
tinct species.

In their postembryonic development, all stomatopods seem
to possess a single postlarval stage, at the termination of their

Stomatopod larvae 527

pelagic larval life. The larval-postlarval molt results in dra-
matic structural changes involving a transition toward the char-
acteristic facies of the benthic adult stages (Manning, 1962;
Alikunhi, 1967). In many species, postlarvae can be correlated
with the adult only by rearing them as was done by K. H.
Alikunhi (1967) in India. Bigelow (1931) was able to as-
sociate the postlarvae with the adults of several different spe-
cies of Pseudosquilla in the Indo-West Pacific region without
utilizing rearing techniques. Studies on the postlarvae are
badly needed to provide basic ontogenetic information in the
stomatopods and to allow the specific identification of the post-
larvae of each species. In addition, postlarval characters may
augment or help to clarify concepts of interspecific relation-
ships.

I am indebted to Thomas E. Bowman, Horton H. Hobbs, Jr.,
and Anthony J. Provenzano, Jr., for commenting on various
portions of the manuscript. The illustrations were made by
my wife Lilly with the support of the Research Awards Pro-
gram of the Smithsonian Institution.

All specimens are in the collection of the Division of Crus-
tacea, National Museum of Natural History, Smithsonian In-
stitution.

POSTLARVAL STAGES OF EASTERN PACIFIC
PSEUDOSQUILLOPSIS
Pseudosquillopsis marmorata (Lockington, 1877 )
Squilla marmorata Lockington, 1877, p. 33. [Type-locality: San Diego,
California].
Figure 1

Material: 1 3,29 mm!'; 2 9, 27-28 mm; Bahia Ballenas, Baja Cali-
fornia; 3 May 1888; Albatross— 1 ¢, 25 mm; 4 9, 25-27 mm; Bahia
San Roque, Baja California; 9-10 February 1950; L. McHugh.— 1 92, 27
mm; San Carlos Bay, Gulf of California; electric light hung over side at
night at anchorage; 30 March 1940; E. F. Ricketts.— 1 9, 28 mm; Gulf
of California; University of California; LXXXIII-H1.—1 ¢, 28 mm; same;
LXVII-H1.

Description: TL 25-29 mm; cornea trilobed, inner portion subdivided
into 2 lobes (Fig. la); antennular peduncle 61-75 percent of carapace
length; width of antennal scale 26-36 percent length; rostral plate pen-

1JIn the postlarvae of Pseudosquillopsis, at least, males can be recognized by the
presence of the buds of the copulatory tubes at the base of the third pair of
walking legs.

528 Proceedings of the Biological Society of Washington

TABLE 1. Summary of basic data for postlarvae and juveniles of
P. lessonii and P. marmorata.

Postlarvae Juveniles
Number of specimens Ws 10 1 1
Total length (mm) (TL) 30-32 25-29 35 40
Carapace length (mm) (CL) 5.7-6.0 4.8-5.4 6.7 rage
Corneal index: range 356-375 327-386 — —
mean 365 365 335 335
Antennular peduncle, as
percent CL: range 75 61-75 — —
mean 75 66 87 78
Antennal scale width, as
percent length: range 37-38 26-36 — —
mean 38 33 29 23
Distance between submedian
teeth of telson, as percent
telson width: range 38-46 46-56 — —
mean 42 50 36 30

Submedian denticles of telson 17 o1-05 = il?

tagonal, length and width subequal, apex not extending beyond cornea,
anterolateral angles rounded (Fig. la); carapace lacking carinae; superior
margin of propodus of claw pectinate, dactylus of claw with 3 well-
formed teeth (Fig. 1b); exposed thoracic somites lacking lateral carinae,
lateral processes of sixth and seventh somites rounded laterally, angled
posteriorly (Fig. lc); anterior 4 abdominal somites unarmed, fifth ab-
dominal somite with posterolateral spines; sixth abdominal somite not
carinate, with 2 pairs of posterior spines, intermediates absent (Fig. 1d);
telson with median carina, remainder of carinae absent (Fig. 1d); sub-
median teeth of telson widely separated, 20-25 submedian denticles pres-
ent on inner surface of each submedian tooth (Fig. le); basal prolonga-
tion of uropod produced into 2 spines, outer longer, with smaller third
spine on inner margin, remainder of inner margin smooth (Fig. lf).

Pseudosquillopsis lessonii (Guérin, 1830)

Squilla lessonii Guérin, 1830, pl. 4, fig. 1; 1838, p. 40 [S. cerisii in text]
[Type-locality: Peru].
Figure 2

Material: 1 &, 32 mm; 1 9, ca. 30 mm; Punta Carretas, Peru; in
stomachs of Neothunnus macropterus, Katsuwonus pelamis, Sarda; En-
rique M. del Solar.

Description: TL 30-32 mm; cornea trilobed, inner portion subdivided
into 2 lobes (Fig. 2a); antennular peduncle 75 percent of carapace

Stomatopod larvae 529

FicurE 2. Pseudosquillopsis lessonii (Guérin), male postlarva, TL 32
mm: d, anterior portion of body; b, raptorial claw; c, lateral processes of
sixth and seventh thoracic somites; d, sixth abdominal somite, telson, and
uropod; e, submedian denticles of telson; f, basal prolongation of uropod.
(Setae omitted ).

length; width of antennal scale 37-38 percent length; rostral plate pen-
tagonal, length and width subequal, apical spine not extending beyond
cormea, anterolateral angles rounded (Fig. 2a); carapace lacking carinae;
superior margin of propodus of claw pectinate, dactylus of claw with 3
well-formed teeth (Fig. 2b); exposed thoracic somites lacking carinae,
lateral processes of sixth and seventh somites rounded (Fig. 2c); anterior
4 abdominal somites unarmed posterolaterally, fifth somite with postero-
lateral spines; sixth abdominal somite with 2 pairs of posterior spines, in-
termediates absent; posterolateral spinule present (Fig. 2d); telson with
single median carina, submedian teeth of telson widely separate, 17 sub-
median denticles present on inner surface of each submedian tooth (Fig.
2e); basal prolongation of uropod produced into 2 spines, outer longer,
with smaller third spine on inner margin, remainder of inner margin
smooth ( Fig. 2f).

COMPARISON OF POSTLARVAL STAGES

The postlarvae of both P. marmorata and P. lessonii can be recognized
as postlarvae by the form of the cornea, for the cornea is trilobed in that

530 Proceedings of the Biological Society of Washington

FicursE 3. Pseudosquillopsis marmorata (Lockington), juvenile female,
TL 40 mm: a, anterior portion of body; b, raptorial claw; c, lateral proc-
esses of exposed thoracic somites; d, sixth abdominal somite, telson, and
uropod; e, submedian denticles of telson; f, basal prolongation of uropod.
(Setae omitted ).

stage of both species. The cornea (Figs. la, 2a) is divided into inner and
outer half by a longitudinal line of cells, and the inner half is further sub-
divided into two lobes. The shape of the eye is characteristic of the
postlarval stage in this genus.

The postlarvae can be identified with Pseudosquillopsis by the pen-
tagonal rostral plate, ornamented anteriorly with a long apical median
projection, the presence of pectinations on the propodus and three well-
formed teeth on the dactylus of the raptorial claw (Figs. 1b, 2b), and the
form of the basal prolongation of the uropod which terminates in two dis-
tal spines with a smaller spine on its inner margin (Figs. lf, 2f).

Stomatopod larvae 531

The postlarvae of species of Pseudosquilla differ from those of Pseudo-
squillopsis in having a slenderer raptorial claw with the propodus lacking
pectinations and dactylus unarmed (the monodactyla stage) and in
having the basal prolongation of the uropod terminate in two spines with
no additional spines on the inner margin. The postlarvae of species of
Parasquilla, which as adults resemble Pseudosquillopsis in many features
(Manning, 1963), differ from those of Pseudosquillopsis in having a
short, rounded rostral plate.

As in the postlarvae of many other species of gonodactylids, the telson
in Pseudosquillopsis postlarvae lacks most of the dorsal ornamentation
characteristic of adults; only the median carina is present. The carination
and spination of the sixth abdominal somite are similarly reduced, for
in the postlarvae only the submedian and the lateral spines are present,
the intermediates being absent, and the spines that are present are not
mounted on carinate ridges.

The postlarvae of P. marmorata may be distinguished from those of P.
lessonii by several features. First, the postlarvae of P. marmorata are
smaller than those of P. lessonii; the 10 specimens of marmorata available
for study range in total length from 25 to 29 mm, whereas the two speci-
mens of lessonii examined possess total lengths of 30 and 32 mm. The
carapace lengths of the postlarvae of marmorata range form 4.8 to 5.4
mm, whereas the carapaces of the two specimens of lessonii measure 5.7
and 6.0 mm. The antennal scale of marmorata is slenderer than that of
lessonii, for in the former species the width of the scale is 26-36 percent
of the length, whereas in lessonii it is 37-38 percent of the length. The
apices of the submedian teeth of the telson are further apart in marmorata
than lessonii; in marmorata the distance between the submedian teeth
ranges from 46—56 percent (mean 50 percent) of the telson width, whereas
in lessonii the distance between the submedian teeth is 38 and 46 per-
cent (mean 42 percent) of the telson width.

The best feature for distinguishing postlarvae of the two species is the
shape of the lateral process of the sixth and seventh thoracic somites. In
marmorata (Fig. 1c) these processes are flattened laterally and angled
posterolaterally, whereas in lessonii (Fig. 2c) they are broadly rounded
laterally and posterolaterally.

JUVENILE STAGES OF EASTERN PACIFIC PSEUDOSQUILLOPSIS

Pseudosquillopsis marmorata (Lockington )
Figure 3

Material: 1 2, 40 mm; La Plata Island, Ecuador; sand, shale, rock, in
45-55. fms; dredge; Hancock Galapagos Expedition, Sta. 212-34; 10
February 1934.

Description: Cornea bilobed, outer margin of eye longer than inner
(Fig. 3a); rostral plate pentagonal, elongate, apical spine not extending
beyond cornea, anterolateral angles acute but not sharp (Fig. 3a); cara-
pace with marginal carinae on posterior fourth; superior margin of prop-

532 Proceedings of the Biological Society of Washington

Ficure 4. Pseudosquillopsis lessonii (Guérin), juvenile male, TL 35
mm: 4, anterior portion of body; b, raptorial claw; c, lateral processes of
exposed thoracic somites; d, sixth abdominal somite, telson, and uropod;
e, submedian denticles of telson (damaged); f, basal prolongation of uro-
pod. (Setae omitted).

odus of claw pectinated, dactylus with 3 well-formed teeth (Fig. 3b);
sixth and seventh thoracic somites with lateral carina, lateral processes
produced into posterior spine (Fig. 3c); fifth abdominal somite with
posterolateral spines; sixth abdominal somite with 3 pairs of spined cari-
nae (Fig. 3d); telson with median carina and 5 pairs of dorsal carinae,
submedians extending onto submedian teeth (Fig. 3d); submedian teeth
of telson separate, submedian denticles present 17 (Fig. 3e); basal pro-
longation of uropod produced into 2 spines, outer longer, with smaller
third spine on inner margin, remainder of inner margin smooth (Fig. 3f).

Stomatopod larvae 533

Pseudosquillopsis lessonii (Guérin)
Figure 4

Material: 1 6, 35 mm; bight on south side of San Juan Bay, Peru;
15°20’S, 75°10’W; bottom dredge; 21 March 1941; M. J. Lobell.

Description: Cornea bilobed, outer margin of eye longer than inner
(Fig. 4a); rostral plate pentagonal, length and width subequal, apical
spine not extending beyond cornea, anterolateral angles rounded (Fig.
4a); carapace with marginal carinae on posterior fourth; superior margin
of propodus of claw pectinate, dactylus with 3 well-formed teeth (Fig.
4b); lateral processes of sixth and seventh thoracic somites lacking carinae
at TL 35 mm, lateral processes rounded posterolaterally (Fig. 4c); fifth
abdominal somite with posterolateral spine; sixth abdominal somite with
3 pairs of spined carinae (Fig. 4d); telson with median carina and 4 pairs
of dorsal carinae, laterals absent at TL 35 mm, submedians interrupted,
not extending onto submedian teeth (Fig. 4d); submedian teeth of telson
separate, submedian denticles present (damaged in available specimen);
basal prolongation of uropod produced into 2 spines, outer longer, with
smaller third spine on inner margin, remainder of inner margin smooth
(Fig. 4f).

CoMPARISON OF JUVENILE STAGES

The juveniles of eastern Pacific Pseudosquillopsis species have assumed
most of the characters of adults. The cornea is bilobed, with the outer
margin of the eye longer than the inner (Figs. 3a, 4a); there is no trace
of the third portion of the cornea found in postlarvae. The rostral plate is
similar to that of adults, but in the two eastern Pacific species, the an-
terolateral angles of the plate in juveniles are unarmed (Figs. 3a, 4a).
The marginal carina is present on the carapace; apparently it is well-
formed even in the first juvenile stage. The carinae and spines of the
sixth abdominal somite are well-developed, although the carinae are not
so strong as in adults. Most of the characteristic carinae of the telson
(median and five pairs) are present in the juvenile stage of marmorata,
TL 40 mn, but in the juvenile of lessonii examined, TL 35 mm, the lateral
carinae are not developed; the juvenile of lessonii also lacks the thoracic
carinae which are clearly developed in the larger specimen of marmorata.

The major difference between the young stages and adults are the pres-
ence of small submedian denticles, 17 in marmorata and an indeterminate
number in lessonii, between the widely separate submedian teeth of the
telson. In adults of Pseudosquillopsis the submedian teeth are appressed
basally and the submedian denticles are completely absent.

The juvenile of P. marmorata, even at TL 40 mm, the smallest speci-
men examined, show the posterolateral spines on the lateral processes of
the sixth and seventh thoracic somites (Fig. 3c). This character will dis-
tinguish this species from the other species in the genus at all sizes beyond
the postlarval stage.

The antennular peduncle is not so long in juveniles of P. lessonii, TL 35
mm, as it is in adults, TL 70 or more.

534 Proceedings of the Biological Society of Washington

TABLE 2. Characters of different stages of Pseudosquillopsis.

Postlarvae Juveniles Adults
Size (mm) 25-33 20-50 70
Cornea Trilobed Bilobed Bilobed
Rostral plate Triangular, with Same Spined laterally
long apical spine; in 2 species
rounded laterally
Carinae on carapace None Reflected marginals Reflected marginals
Claw 3 teeth, propodus Same Same
pectinate
Sixth abdominal 2 pairs of non- 3 pairs of Same
somite carinate spines carinate spines
Carinae on telson Median Median and Median and 5 pairs
4-5 pairs
Submedian denticles Present Present Absent
of telson
Basal prolongation Long outer spine, Same Same
of uropod with 2 smaller
spines on inner
margin

The two available juveniles are not of comparable age, as evidenced by
the better development of carination in the specimen of marmorata at TL
40 mm than in the specimen of lessonii at TL 35 mm. Both specimens,
however, are clearly subadults, and as such have been used to show transi-
tion in development between the postlarva and the adult. Rearing ex-
periments obviously are needed to provide more detailed information on
changes in postlarval development and the stages and sizes at which they
occur.

GENERAL DISCUSSION

The two postlarvae from the Gulf of Guinea which prompted this study
clearly can be identified as the postlarvae of Pseudosquillopsis. They are
tentatively identified with P. cerisii (Roux, 1828), the only species of the
genus known from the eastern Atlantic region. The postlarvae of P.
cerisii, TL 30-33 mm, are very similar to those of P. lessonii, TL 30-32
mm. However, the apex of the rostral plate in the postlarva of P. cerisii
is shorter and blunter than that of the postlarva of P. lessonii, and the dis-
tance between the apices of the submedian teeth of the telson is greater
in the postlarva of P. cerisii than in that of P. lessonii.

Characters of postlarvae, juveniles, and adults of members of the
genus Pseudosquillopsis are summarized in Table 2. The early stages are
distinctive but are clearly referable to Pseudosquillopsis, which was here-
tofore based on characters afforded by adults only. The most important
difference between young specimens and adults is the presence of sub-
median denticles on the telson in the former and their absence in the adult
stage.

Young specimens, possibly including both postlarvae and juveniles, of

Stomatopod larvae 535

P. dofleini (Balss) from Japan have been recorded by Komai (1927). He
noted that in the smallest specimen, TL 20 mm, the inner margin of the
basal prolongation was unarmed. He also noted that the eyes of his small
specimens were subsimilar to those adults. In view of the latter observa-
tion, his specimens were probably juveniles rather than postlarvae. From
the length of the smallest specimens recorded by him, TL 20 mn, it
might be assumed that the postlarvae of P. dofleini are smaller than those
of the remaining species of the genus.

Adults of the two Eastern Pacific species of the genus, which have the
anterolateral angles of the rostral plate armed with spines and also have
a smooth inner proximal margin on the basal prolongation of the uropod,
are more closely related to each other than to either the Atlantic species
or the one found in the Indo-West Pacific region. Indeed, the extra-
American species, P. cerisii and P. dofleini, both of which have spinules
proximally on the basal prolongation of the uropod, are extremely difficult
to separate.

In his description of P. dofleini, Balss (1910) noted that it differed
from P. cerisii in having the basal prolongation of the uropod armed with
spinules; small spinules are definitely present on the basal prolongation in
the only specimen of P. cerisii I have examined, a male, TL 87 mm, from
Naples, Italy, and these spinules have been noted by other authors as
well (see Seréne, 1962, fig. 2c). The spinules in P. dofleini apparently
differ from those found in P. cerisii in that they increase in size distally,
with the distalmost not markedly smaller than the innermost of the three
terminal spines on the basal prolongation, whereas all of the spinules are
small in P. cerisii. In other respects P. dofleini and P. cerisii resemble
each other very closely.

The rostral plate is unarmed anterolaterally in the single specimen of
P. cerisii which I have examined. Seréne (1962, p. 16), in his diagnosis of
Pseudosquillopsis, stated that the anterolateral angles were armed in P.
cerisii and P. lessonii, but the plate is rounded anterolaterally in the speci-
men I examined.

The species placed in Pseudosquillopsis share the following features as
adults: cornea bilobed, with outer margin of eye longer than inner; cara-
pace lacking cervical groove, ornamented with short marginal carinae
only; propodus of raptorial claw stout, superior margin lined with numer-
ous short, blunt projections; dactylus of claw armed with three teeth;
dorsal surface of telson ornamented with median carina and 5 pairs of
dorsal carinae; submedian teeth of telson with bases appressed, sub-
median denticles completely suppressed; basal prolongation of uropod
terminating in 2 spines, outer larger, with smaller third spine on inner
margin. More detailed diagnoses have been provided by Seréne (1962)
and Manning (1963), both of whom also commented on the relationships
of Pseudosquillopsis to Parasquilla and Pseudosquilla.

The provisional key to the species of Pseudosquillopsis presented below
may have to be revised when more material of P. dofleini is available for
study.

536 Proceedings of the Biological Society of Washington

PROVISIONAL KEY TO ADULTS OF PSEUDOSQUILLOPSIS
Ly Inner half of basal prolongation of uropod with spinules; rostral

plate rounded laterally 2,
Inner half of basal prolongation of uropod smooth or with low,
rounded tubercles; rostral plate with lateral spines 3

2. (1)Spinules on inner margin of basal prolongation of uropod in-
creasing in size distally, inner spine of basal prolongation not
markedly larger than distalmost spinule —-__-_----______-
eee ee eee _ P. dofleini (Balss, 1910): Japan.

Spinules on inner margin of basal prolongation of uropod small,
not markedly increasing in size distally, outermost much
smaller than inner spine 2 ee P
cerisii (Roux, 1828): Mediterranean Sea, Gulf of Guinea

3. (1)Antennular peduncle as long as or longer than carapace; lateral
processes of sixth and seventh thoracic somites rounded pos-
terolaterally __.____ P. lessonii (Guérin, 1830): Peru, Chile,
Juan Fernandez Island.

Antennular peduncle shorter than carapace; lateral processes of
sixth and seventh thoracic somites spined posterolaterally —_
P. marmorata (Lockington, 1877): southern California, Gulf
of California, Galapagos Islands.

LITERATURE CITED

Auikunut, K. H., 1967. An account of the post-larval development,
moulting and growth of the common stomatopods of the
Madras coast, pp. 824-939, figs. 1-94, pls. 1-3. In Proceed-
ings of the Symposium on Crustacea, Marine Biol. Assoc. In-
dia, pt. II: iv + 945 pp.

Bass, H., 1910. Ostasiatische Stomatopoden. Beitrage zur Naturge-
schichte Ostasiens, herausgegeben von Dr. F. Doflein. Abh.
math.-phys. Klasse Bayer. Akad. Wiss. Miinchen, Suppl.-Bd.
Il, 2: 1-11, figs. 1-2.

BicELow, R. P., 1894. Report on the Crustacea of the Order Stomatopoda
collected by the Steamer Albatross between 1885 and 1891,
and on other specimens in the U.S. National Museum. Proc.
U.S. Nat. Mus., 17 (1017): 489-550, figs. 1-28, pls. 20-22.

, 1931. Stomatopoda of the southern and eastern Pacific Ocean
and the Hawaiian Islands. Bull. Mus. Comp. Zool., Harvard,
72(4): 105-191, figs. 1-10, pls. 1-2.

GiesprEcHT, W., 1910. Stomatopoda. Fauna u. Flora. Neapel, monogr.
33: vii + 239, pls. 1-11.

GuERIN-MENEVILLE, F. E., 1829-1830. Crustacés; pls. 1-5. In Duper-
rey, L. I., Voyage autour du monde, exécutés par ordre du
Roi, sur la corvette de sa Majesté, La Coquille, pendant les
années 1822, 1823, 1824, et 1825. Atlas, pls. 1-24. Paris,
Arthus Bertrand [see Holthuis, L. B., 1961, Crustaceana, 3
(2): 168, for dates of publication of crustacean plates].

Stomatopod larvae 537

———., 1838. Crustacés, Arachnides et Insectes, pp. xiv, 9-47. In
Duperrey, L. I., Voyage autour du monde. . ., Zoologie, 2(2)
1: xiv + 319. Paris, Arthus Bertrand.

Komat, T., 1927. Stomatopoda of Japan and adjacent localities. Mem.
Coll. Sci. Kyoto Imp. Univ., (B) 3(3): 307-354, figs. 1-2,
pls. 13-14.

Locxincrton, W. N., 1877. Remarks on the Crustacea of the Pacific coast,
with descriptions of some new species. Proc. California Acad.
Sci., 7: 28-36 [pp. 1-9 on separate].

Manninec, R. B., 1962. Alima hyalina Leach, the pelagic larva of the
stomatopod crustacean Squilla alba Bigelow. Bull. Mar. Sci.
Gulf & Carib., 12(3): 496-507, figs. 1-4.

, 1963. Preliminary revision of the genera Pseudosquilla and
Lysiosquilla with descriptions of six new genera (Crustacea:
Stomatopoda). Bull. Mar. Sci. Gulf & Carib., 13(2): 308—
328.

Miers, E. J., 1880. On the Squillidae. Ann. Mag. Nat. Hist., 5 (5): 1-30,
108-127, pls. 1-3.

MiILNE-Epwarps, H., 1837. Histoire naturelle des Crustacés, compren-
ant l’anatomie, la physiologie et la classification de ces ani-
maux, 2:1-532. Atlas: 1-32, pls. 142. Paris, Roret.

Roux, P., 1828-1830. Crustacés de la Méditerranée et de son littoral: iv
176, pls. 1-45. Paris; Marseille.

Scumitr, W. L., 1926. The macruran, anomuran, and stomatopod crus-
taceans collected by the American Museum Congo Expedi-
tion, 1909-1915. Bull. Amer. Mus. Nat. Hist., 53: 1-67, figs.
1-75, pls. 1-9.

, 1940. The stomatopods of the west coast of America, based
on collections made by the Allan Hancock Expeditions, 1933-
1938. Allan Hancock Pacific Expeds., 5(4): 129-225, figs.
1-33.

SERENE, R., 1962. Révision du genre Pseudosquilla (Stomatopoda) et
définition de genres nouveaux. Bull. Inst. océanogr. Monaco,
no. 1241: 1-27, figs. 1-5.

538 Proceedings of the Biological Society of Washington

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558 17 November 1969

PROCEEDINGS
OF THE

BIOLOGICAL SOCIETY OF WASHINGTON

FRESHWATER TRICLADS (TURBELLARIA ) OF
NORTH AMERICA. I. THE GENUS PLANARIA.)

By RoMAN KENK
Senior Scientist, The George Washington University, and
Research Associate, Smithsonian Institution

The genus Planaria was established by O. F. Miiller (1776:
221) to separate the free-living lower worms from the parasitic
trematodes which retained the older name Fasciola. Origi-
nally Planaria comprised all known Turbellaria living in fresh
water, in the sea, and on land and, besides these, the present
Nemertina or Rhynchocoela. The extent of the genus was
gradually narrowed as newly established genera were separated
from it, chiefly by Dugés (1528), Ehrenberg (1831), and
Orsted (1843 and 1844). After Ehrenberg’s revision of the
system, which first introduced the name Turbellaria, Planaria
was restricted to turbellarians with branched intestine (“Den-
drocoela” ) which possessed two eyes. Orsted, who further re-
fined the systematic arrangement of the “flatworms,” separated
the polyclads (“Cryptocoela”) from the Dendrocoela and ap-
plied the name Planaria mainly to triclads, both freshwater
and marine, including also the many-eyed species which
Ehrenberg had separated from Planaria. In 1844 (p. 51) he re-
moved from it the new genus Dendrocoelum on the basis of its
intestinal branching. In the following years the name Planaria
was used rather indiscriminately for many turbellarian forms.
With the progress of the studies of the internal structure of the
various turbellarian taxa in the second half of the nineteenth
century it gradually became restricted to freshwater triclads.

1 Supported by National Science Foundation Grant GB-6016 to the George Wash-
ington University.

2 Author’s address: Department of Invertebrate Zoology, Museum of Natural
History, Smithsonian Institution, Washington, D. C. 20560.

46—Proc. Biot. Soc. WaAsH., Vou. 82, 1969 (539)

540 Proceedings of the Biological Society of Washington

Even today the common name “planarian” may signify any
triclad species (freshwater, land, and sea planarians for Pal-
udicola, Terricola, and Maricola ).

It was not until the twentieth century that further important
studies were reflected in the classification of the freshwater
triclads. Thus Komarek (1926) proposed a more “natural”
system of the Paludicola for the European representatives of
this group. He established the genera Dendroplanaria (for
Miiller’s Fasciola torva), Fonticola, and Albiplanaria, revived
Hesse’s (1897) Euplanaria, and restricted the generic name
Planaria to Dana’s (1766) Hirudo alpina.

The present system of the freshwater triclads, which is in
principle being followed by modem authors, was introduced
by Kenk (1930). The suborder Paludicola or Probursalia of the
order Tricladida is divided into two families, the Planariidae and
the Dendrocoelidae. A third family, Kenkiidae, split off from the
Planariidae by Hyman (1937) appears not to be justified and
has not been accepted by some later planarian workers (e.g., de
Beauchamp, 1961: 103; Mitchell, 1968: 615-61S ).

The two families are distinguished by the arrangement of
the muscle fibers in the internal muscular zone of the pharynx,
which in the Planariidae consists of two distinct layers, a layer
of circular fibers adjoining the internal epithelium, followed by
a layer of longitudinal fibers; in the Dendrocoelidae the in-
ternal muscular zone is represented by a single layer of inter-
mingled circular and longitudinal muscle fibers. Fortunately,
this characteristic can be recognized even in sexually immature
specimens, while many of the generic and specific features
used in the identification of the smaller taxa concern primarily
the reproductive system.

Komarek’s (1926) restriction of the genus Planaria to P. al-
pina and its immediate relatives was untenable, since the spe-
cies alpina was not included among the species assigned to the
genus Planaria when it was first established by O. F. Miller
(1776). Kenk (1930: 293) therefore selected Fasciola torva
Miller (1774) as the type of the genus and presented a defini-
tion of the genus, slightly emended in a later paper (1935:
111): Planariidae whose oviducts—without embracing the

The genus Planaria 541

stalk of the bursa copulatrix (or forming a loop around it)—
unite to a common oviduct which opens into the genital atrium.
Male atrium without radial muscle plates. Adenodacty] present,
constructed according to the Planaria torva type. (An analysis
of the adenodactyl was given by Kenk, 1930: 159. )

In this narrower sense the genus Planaria comprises very few
species, scattered over three continents: P. torva (Miller,
1774) with its probable synonym P. onegensis Zabusov (1901),
in Europe and possibly Asia; P. kempi Whitehouse (1913), in
India; and P. dactyligera Kenk (1935), in North America.

A detailed review of the present status of the type species,
Planaria torva, has been presented recently by Ball, Reynold-
son and Warwick (1969).

Planaria dactyligera dactyligera Kenk, 1935

Type material: Holotype, from Mountain Lake, Giles County, Virginia,
2 slides of sagittal sections, U. S. National Museum No. 39461. Paratypes
in the author’s collection.

The species Planaria dactyligera has been described in detail in an
earlier paper (Kenk, 1935: 105-110) from material collected in several
localities in Virginia. Examination of specimens from North Carolina made
it advisable to distinguish two subspecies of this species. The principal
characteristics of the typical form from Virginia may be briefly recapitu-
lated here.

External features: Mature animals are up to 13 mm long and 1.75 mm
wide. The dorsal side is darkly pigmented, gray, brown, or black; the
ventral surface, somewhat lighter. The anterior end is truncate, with al-
most straight frontal margin and rounded lateral (auricular) edges. In
the quietly gliding animal there may be an insignificant narrowing or neck
behind the auricles. Eyes are normally two, placed rather close together
(about 4% the width of the head at the level of the eyes) and removed
from the frontal margin by a distance slightly less than the width of the
head.

Reproductive system: The main features distinguishing the species
from its relatives are in the anatomy of the reproductive system (cf.
Kenk, 1935, figs. 25 and 27). The testes are predominantly ventral and
occupy a pair of broad bands, one on either side of the midline, extend-
ing from a short distance behind the eyes to about the level of the mouth
opening. The vasa deferentia expand in the region of the pharynx to
form a pair of sinuous spermiducal vesicles (or false seminal vesicles ),
filled with sperm, as is typical for freshwater triclads in general. They ap-
proach the bulb of the penis from the antero-lateral sides. The two ovi-
ducts (or ovovitelline ducts), which in their main course accompany the

542 Proceedings of the Biological Society of Washington

ventral nerve cords, turn upward and medially in the region of the copula-
tory complex and unite in the space above the male atrium and below the
stalk of the copulatory bursa to form the common oviduct.

There is no distinct common genital atrium developed, as the various
cavities of the copulatory organs meet in the immediate vicinity of the
genital aperture: from the anterior side the male atrium, dorsally the
duct of the copulatory bursa, and posteriorly the opening of the adeno-
dactyl. The male atrium is more or less cone-shaped, duplicating the
shape of the penis which it encloses.

The penis consists of a spherical bulb embedded in the mesenchyme
and a conical papilla protruding into the male atrium. At the transition
between the two parts is a cavity (the shape of which may vary according
to the contraction or expansion of the organ), the seminal vesicle. This
cavity receives from its anterior side the two vasa deferentia which have
entered the bulb from the sides and have formed a few convolutions within
the bulb, with a common opening. From the seminal vesicle the ejacula-
tory duct emerges as a tapering, straight canal which opens at the tip of
the papilla. Many gland ducts penetrate the penis bulb from the sur-
rounding mesenchyme and open into the seminal vesicle.

The muscular coat underlying the outer epithelium of the penis papilla
consists of two layers: a circular layer adjoining the epithelium, followed
by a layer of longitudinal fibers. The thickness of this muscle coat is
about equal to, or slightly greater than, that of the wall of the male atrium.

The common oviduct formed by the union of the paired oviducts enters
the posterior part of the male atrium from the dorsal side or from the left.
The terminal parts of the paired oviducts and almost the entire common
oviduct receive very numerous gland ducts filled with an intensely
eosinophilic secretion.

The copulatory bursa is a large sac lying between the pharyngeal
pouch and the bulb of the penis. Its duct or stalk, running dorsally
to the male atrium, curves postero-ventrally and joins the atrial com-
plex close to the gonopore. There is no distinct posterior portion or
vagina developed.

The adenodactyl is a large hollow organ lying behind the genital
aperture and opening close to the aperture without a prominent protrud-
ing papilla. Its heavy muscular wall is pierced by numerous gland outlets
emptying its inner cavity.

Distribution: The type locality of Planaria dactyligera dactyligera
is Mountain Lake, Giles County, Virginia, near south bank of the lake.
It has been collected also in Rockbridge, Highland, and Albemarle coun-
ties, Virginia (Kenk, 1935: 109). Fite (1952), who studied a nematode
parasitic in the pharynx of this species, collected his material at Twin
Springs near the Mountain Lake Biological Station of the University of
Virginia, in Giles County.

Chandler (1966: 11) reports that he collected some, mostly immature,
planarians near Bloomington, Monroe County, Indiana, which he tenta-
tively identified as Planaria dactyligera. He kindly sent me a slide of the

The genus Planaria 543

problematic species for examination. The preliminary identification
proved to be erroneous.

Planaria dactyligera musculosa new subspecies®

Type material: Holotype, from Ossipee, Alamance County, North
Carolina, 4 slides of sagittal sections, U. S. National Museum No. 39462.
Paratypes, 13 series of sagittal and transversal sections, in the author’s
collection.

External features: Mature specimens attain a length up to 11 mm and
a width of 1.3 mm. The head is truncate, with the frontal margin slightly
bulging in its entire extent (Fig. la) or in the central portion (Fig. 1b).
In quiet gliding either of these two shapes may be assumed transitorily.
The lateral edges are rounded. A very slight narrowing (neck) may be
seen behind the head, then the body margins gradually diverge, remain
parallel for some distance, converge again behind the pharyngeal region,
and meet in the bluntly pointed posterior end. There are two eyes, lying
close together (less than 4% the width of the head at the level of the
eyes) and farther distant from the frontal margin than from the lateral
margins.

The pigmentation of the dorsal surface is usually dark, almost black,
appearing somewhat cloudy under magnification. Only the two eye
patches are free of pigment. There are, however, lighter areas visible
above the pharynx and, in sexually mature specimens, above the copula-
tory complex. The ventral side is also pigmented, but in a lighter hue
than the dorsal side. The mouth opening is visible as a distinct white
spot, the gonopore is not quite as clearly discernible. Freshly hatched
young are unpigmented, white, and acquire their pigmentation gradually
during their gowth and development. Animals kept in cultures in the dark
tend to be more lightly pigmented than specimens in their natural habitat.

The pharynx is inserted at about the middle of the body, its length
being approximately 4; the body length. The mouth and the gonopore
divide the posterior half of the body into three almost equal thirds.

As is seen from this description, Planaria dactyligera musculosa cannot
be distinguished from P. d. dactyligera by its external features. It also re-
sembles closely some pigmented species of Phagocata (P. velata [Stringer],
P. vernalis Kenk, P. crenophila Carpenter, and at least one other,
undescribed, species of this genus) as well as Hymanella retenuova Castle.

Internal characters: The anatomical characteristics of Planaria dactyli-
gera musculosa conform in most particulars with those of the typical form
(cf. Kenk, 1935: 105-109). The body pigmentation obscures the intes-
tinal branching in the living specimens. The anterior end of the pre-
pharyngeal intestinal ramus extends far into the head, forming a straight,
unbranched diverticulum which reaches to a level anterior to the eyes

(Fig. 1b).

3 musculosus, Latin, muscular, referring to the circular muscle layer of the penis
papilla.

544 Proceedings of the Biological Society of Washington

cee

5mm

The genus Planaria 545

The testes are predominantly ventral, but individual follicles may be
displaced dorsally or, at full maturity, may occupy the entire dorsoven-
tral diameter of the body. The testicular zone of each side extends from a
level behind the eyes to approximately the level of the mouth opening.
The ovaries (with large parovaries), vitellaria, ovovitelline ducts, and
vasa deferentia do not deviate from the conditions seen in the typical
form. There are, however, distinct differences in the copulatory apparatus
of the two subspecies.

A semidiagrammatic view of the copulatory apparatus of Planaria
dactyligera musculosa, with particular reference to its muscular and
glandular differentiations, is shown in Figure 4. In comparing this figure
with the corresponding diagram for P. dactyligera dactyligera (Kenk,
1935, Fig. 27), the differences in the proportions of the individual organs
should be disregarded, as the present figure is based on an unusually well
extended specimen while that of the type-species shows a certain amount
of longitudinal contraction such as is commonly encountered in preserved
planarians.

In the new subspecies, the large copulatory bursa (b) regularly shows
numerous lobes or diverticula projecting mainly in the lateral direction.
The bursa duct (bd) gradually widens as it curves toward the gonopore,
but shows no histologically distinct or sharply demarcated vagina. It
opens into the atrial complex close to the gonopore (gp).

The penis consists of a spherical bulb containing loosely arranged mus-
cle fibers running in more or less concentric layers, and the end parts of
the sinuous and highly muscular vasa deferentia (vd). The bulb is
pierced by numerous gland ducts which enter it from a wide area of the
surrounding mesenchyme and open into the seminal vesicle (vs).

The two vasa deferentia empty, by a common opening, into the cavity
of the penis which shows an anterior wider part (seminal vesicle, vs)
and tapers posteriorly to a narrower ejaculatory duct which opens at the
tip of the papilla. One of the chief distinguishing features of the new
subspecies is the extraordinary development of the external circular
muscle layer (mp) at the basis of the penis papilla. The thickness of this
layer is several times the thickness of the muscle layers of the male atrium
(while in P. dactyligera dactyligera the corresponding muscle layers are
about equal in thickness ).

<

Fic. 1-3. 1. Planaria dactyligera musculosa. a. Quietly gliding animal;
gp, gonopore; m, mouth; ph, pharynx. b. Anterior end, showing position
of eyes and intestine. c. Freshly hatched young; the shaded area indicates
the extent of the intestinal trunks and branches. 2. Planaria dactyligera
musculosa, photograph from life, «9. 3. Planaria occulta, photograph
from life, x9.

‘QPISAA [RUIUAS ‘sa ‘SUdTOJEpP SPA “pa ‘oNnpPIAO UOUULOD ‘po
serpided stuad Jo IAL] BpPOSNUT Aepno.t1o “duu syynour “Ud ‘atodouos ‘da ‘ypeys Bsinq “pq ‘esinq Aioyepndoo “q ‘[AJovpouope “pp ‘[Ayorp
-ouape JO WUNEQR ‘pp “UOTOoS [eyes UL sursso0 ALOy[NAOD Jo MIA OVULLUBISvIPIUlas “DsOpNISNUL piaayAyoop Diavud]g =" F “OW

po po d6 pq po SA dw pA q wi

ty of Washington

1e

Wu |

546 Proceedings of the Biological Soc

The genus Planaria 547

The second difference between the two forms concerns the relation of
the adenodactyl (ad) to the atrial complex. The adenodactyl is a large
ovoid or pear-shaped organ with a thick muscle coat of chiefly circular
fibers, very densely arranged, with the corresponding cell nuclei forming
a peripheral layer. Its elongated cavity opens into a separate part of the
genital atrium (aa) which narrows anteriorly and connects with the
general atrial complex very close to the genital opening (gp). There is no
distinct papilla of the adenodactyl protruding into this chamber. The
epithelium of the chamber is pierced by very many gland outlets
originating in the mesenchyme.

In summary, the main distinguishing characteristics of the new sub-
species are (1) the thick circular muscle layer at the base of the penis
papilla and (2) the presence of a highly gandular antechamber between
the gonopore and the adenodactyl.

Distribution: The first specimens of Planaria dactyligera musculosa
were sent to me by Dr. T. E. Powell, Jr., of the Carolina Biological Supply
Company. They had been collected at the “Sawdust Pile Location” in
Ossipee, Alamance County, North Carolina, in November 1966. The
majority of the animals were sexually mature.

Additional specimens were collected on 13 June 1968 in two localities
on the grounds of the Warren Laboratories of the Carolina Biological
Supply Company, off U.S. Highway 158, 3 miles E of Warrenton,
Warren County, North Carolina. One was a stream near the side en-
trance road to the Laboratories, the other on the airport road. The speci-
mens were small, immature, but matured in the laboratory cultures.

Reynierse and Ellis (1967) report that they used Planaria dactyligera
in an experiment on planarian behavior. Since they had obtained their
animals from the Carolina Biological Supply Company (personal com-
munication ), we may safely assume that their planarians belonged to the
subspecies P. d. musculosa.

Longest (1966: 39-41) reported Planaria dactyligera from Abita
Springs State Park, St. Tammany Parish, Louisiana. Examination of his
slides showed that his specimens belonged to the subspecies musculosa.

Parasites: Small nematodes were occasionally observed in the mesen-
chymatous zone of the pharynx. They apparently were capable of moving
freely through the tissue as no cysts were formed around them. The
holotrichous ciliate, Sieboldiellina planariarum (Siebold) occurred in
the lumen of the intestine and sometimes in the pharyngeal pouch. Some
of the specimens were infested with the peritrichous ciliate epizoite, Ur-
ceolaria mitra (Siebold), attached mainly to their dorsal surfaces.

Observations in laboratory cultures: Cultures of Planaria dactyligera
musculosa were kept in spring water in a constant-temperature chamber
at about 14°C, and fed beef liver and/or Tubifex which were readily
taken. The worms produced cocoons all year round. The cocoons are
ellipsoidal, rather variable in size, the longest diameter measuring 0.8—
1.7 mm, the shortest 0.6-1.0 mm. When deposited they are attached to

548 Proceedings of the Biological Society of Washington

SF ee

ph vd vs de bd ad

Fic. 5-6. 5. Planaria dactyligera musculosa, sagittal section of copula-
tory complex, showing the adenodactyl (ad) with its atrial chamber (aa),
the terminal part of the bursa duct (bd), the thick muscular coat (mp)
of the penis papilla (excentrically cut), and part of the common oviduct
(od), «113. 6. Planaria occulta, sagittal section through copulatory com-
plex, showing the position of the pharynx (ph), vas deferens (vd), sem-
inal vesicle (vs), the convoluted ejaculatory duct (de), and parts of the
bursa duct (bd) and adenodactyl (ad), «113.

The genus Planaria 549

Fic. 7. Planaria occulta, outline drawing of gliding animal, cop, cop-
ulatory complex; ph, pharynx.

the substrate by a colorless jelly-like substance, the long axis being
parallel to the surface of the substrate (bottom or side wall of the aquar-
ium). They easily loosen their attachment when the cultures are handled.
As is the rule in triclad cocoons, the freshly laid egg capsule is of a light
reddish-brown color and darkens in a few days to become dark brown to
almost black. Three to 14 young hatched from a single cocoon after 3-4
weeks. The freshly hatched young (Fig. lc) vary in size from 1.5 to 3
mm in length. They are unpigmented, white, with a rounded head end.
As they grow in size, they gradually become pigmented and acquire the
typical truncate head shape characteristic of older specimens.

No asexual reproduction by fission was observed during 28 months of
culturing.

Planaria occulta new species*

Type material: Holotype, from Duffield, Scott County, Virginia, 2
slides of sagittal sections, U. S. National Museum No. 39463. Paratypes,
sagittal and transversal sections of 7 specimens, in the author’s collection.

4 occultus, Latin, hidden, alluding to the subterranean occurrence in a well.

590 Proceedings of the Biological Society of Washington

a b C

Fic. 8. Anterior ends of three similar planarian species: a, Planaria oc-
culta; b, Phagocata morgani; c, Phagocata oregonensis.

External features (Fig. 3): Mature specimens measure up to 9 mm in
length and 1.5 mm in width when gliding quietly. The species is without
body pigment, appearing white when the intestine contains no colored
matter; even with the intestine filled, the head, the lateral margins of the
body, and the places occupied by the pharynx and copulatory apparatus
are always white. The head is truncated, with a slightly convex frontal
margin and rounded lateral edges (Fig. 8a). In quiet gliding it may tran-
sitorily present a moderately bulging median section (Fig. 7). There is
no neck constriction behind the head. The lateral margins gradually di-
verge, soon become parallel, start converging again in the region of the
pharynx, and meet at the moderately pointed posterior end. There are
two eyes, rather far removed from the frontal margin, their distance from
each other amounting to about one-fourth the width of the head at eye
level. The distance of each eye from the lateral margin is smaller than
that from the frontal margin. No supernumerary eyes, such as frequently
occur in normally two-eyed planarians, have been observed in this spe-
cies. The pharynx is inserted at about the middle of the body and amounts
in length to approximately one-fifth the body length. In sexually mature
specimens, the copulatory complex occupies the anterior three-fourths of
the postpharyngeal region.

At first glance, Planaria occulta resembles the common Phagocata
morgani (Stevens & Boring) with which it shares its geographic area, and
other unpigmented species of Phagocata of North America (P. nivea
Kenk, P. oregonensis Hyman), Europe, and Japan. Unfortunately not all
these species have been adequately described in the living state. In com-
paring specimens in good physiological condition during gliding locomo-
tion, one may discover subtle differences between these various species,
differences which are entirely obscured in the preserved animals. Figure
8 shows such a comparison between P. occulta (a), P. morgani from a
spring in Rock Creek Park in Washington, D. C. (b), and P. oregonensis
from Portland, Oregon (c). It will be noticed that in P. occulta the
anterior intestinal ramus ends at a level anterior to the eyes, while in the

The genus Planaria dol

| | | |

ep nu ma i

Fic. 9-10. Planaria occulta. 9. Paramedian section through the ante-
rior end, showing the auricular sense organ (au) and the marginal adhe-
sive zone (az), X310. 10. Cross section of postpharyngeal region at the
level of the adenodactyl; ep, ventral surface epithelium; i, intestinal epi-
thelium; ma, muscle layer of adenodactyl; nu, layer of muscle cell bodies
and nuclei. «310.

552 Proceedings of the Biological Society of Washington

adult P. morgani and P. oregonensis the intestine is confined to a region
posterior to the eyes. It must be mentioned, however, that freshly
hatched young of P. morgani show an anterior extension of the intestinal
ramus between the eyes similar to that of P. occulta.

The locomotion of Planaria occulta is a smooth gliding. No “crawling”
movements such as are observed in many other species have been seen
even upon mechanical stimulation of the animal (to which they react by a
brief contraction of the body, followed immediately by continued
gliding ).

Integument: The epithelium of the general surface shows no peculi-
arities, the cells of the dorsal epithelium being somewhat taller than those
of the ventral side (about 12 u and 8 uw, respectively, depending somewhat
on the contraction of the body). No distinct adhesive organ is developed.
A narrow band of gland openings runs ventrally along the margins of the
body, the marginal adhesive zone (Figs. 9 & 11, az). This band is inter-
rupted only in the center of the frontal margin of the head by a very short
(30 »-35 w) gap. The secretion of the adhesive glands is granular and
strongly eosinophilic.

Sense organs: In addition to the eyes, there are other sensory structures
discernible in the head region. The auricular sense organs occupy the
lateral parts of the frontal margin and consist of strips of modified epithe-
lium, densely ciliated and containing only few rhabdites which are gen-
erally much shorter than those of the surrounding epithelia. There
are no sensory pits or grooves developed, as the organs form the very
edge of the margin, being separated from the adhesive gland zone by a
narrow band of normal surface epithelium. Another sensory area, a small
patch with similarly modified rhabdite-free epithelium, lies on the ventral
side of the head, immediately behind the gap of the marginal adhesive
zone.

Digestive system: The pharyngeal muscles conform with the typical
arrangement in the family Planariidae, the muscle fibers of the internal
zone forming two separate layers, an inner circular and an outer longi-
tudinal one. The external muscle zone consists likewise of two layers, a
layer of longitudinal fibers underlying the outer epithelial covering, fol-
lowed by a layer of circular fibers. There is no third (longitudinal) layer
developed. The anterior intestinal ramus which, as indicated above, ex-
tends in the head region to a level in front of the eyes, bears on either
side 6 to 9 branches; each posterior ramus, 13 to 19 shorter and less pro-
fusely ramified branches.

Reproductive system: The numerous testes occuply a longitudinal
zone on either side of the midline, each zone extending from a short dis-
tance behind the head to approximately the level of the mouth opening. In
a prepharyngeal cross section (Fig. 11) one may see on each side one to six
more or less rounded testicular follicles (¢), situated in the ventral parts
of the mesenchyme, mainly below the intestinal branches (i) and above
the ventral nerve cords (n). Individual testicles, particularly at full ma-
turity, may penetrate dorsally in the spaces between the branches of the

The genus Planaria 503

m vd vs de am gp ad

Fic. 11-12. Planaria occulta. 11. Transversal section of prepharyn-
geal region. 12. Semidiagrammatic view of copulatory apparatus in sagit-
tal section. ad, adenodactyl; am, male atrium; az, marginal adhesive
zone; b, copulatory bursa; bd, bursa stalk; de, ejaculatory duct; gp, gono-
pore; i, intestine; m, mouth; n, ventral nerve cord; od, oviduct; odc, com-
mon oviduct; t, testis; vd, vas deferens; ve, vas efferens; vi, yolk glands;
vs, seminal vesicle.

intestine. From each testicle a delicate duct, the vas efferens (ve), pro-
ceeds ventrally to open into the likewise very thin-walled sperm duct or
vas deferens (vd) of the corresponding side, which runs along the medial
border of the ventral nerve cord, close to the subcutaneous muscle layer
of the ventral surface. In the region of the pharynx the vasa deferentia
expand greatly in diameter and are seen as a pair of tortuous tubes filled

504 Proceedings of the Biological Society of Washington

with sperm, the false seminal vesicles (or spermiducal vesicles according
to Hyman’s [1951: 113] terminology). At the level of the penis they
bend upward and enter the bulb portion of the penis from the sides.

The rather small spherical ovaries or germaries are in the typical posi-
tion, a short distance behind the eyes, adjoining the medial side of the
ventral nerve cords. Each ovary is accompanied by more voluminous,
lobate masses of cells extending toward the dorsal side, the parovaries.
The cytoplasm of these cells stains dark blue with Ehrlich’s hematoxylin.
The oviducts or ovovitelline ducts start from the lateral surfaces of the
ovaries, each beginning with a slightly widened portion, the seminal re-
ceptacle. They run caudally along the dorso-lateral side of the ventral
nerve cords (Fig. 11, od). On their course they connect with numerous
yolk glands or vitellaria (vi) which, at full maturity, represent volumi-
nous masses occupying chiefly the dorsal and lateral portions of the mes-
enchyme from the level of the ovaries to almost the posterior end of the
body.

The copulatory apparatus (Fig. 12) occupies, in sagittal sections, the
greater part of the postpharyngeal region. The genital aperture or
gonopore (gp) is situated far caudally, its distance from the mouth open-
ing (m) being about twice the distance from the tail end of the body.
There is no distinctly developed common genital cavity or atrium, as the
various ducts of the copulatory complex meet almost at the gonopore:
from the anterior side the male atrium (am), dorsally the duct of the
copulatory bursa (bd), and from the caudal side the outlet of a small
cavity connected with the adenodactyl (ad). The “male” atrium en-
closes the papilla of the penis and receives in its posterior portion the
mouth of the common ovovitelline duct (odc). Its lining is an epithelium
of cubical, ciliated cells below which are two layers of muscles, a layer
of circular fibers and below it one of longitudinal fibers.

The male copulatory organ or penis consists of a moderately developed
spherical bulb embedded in the mesenchyme a short distance behind
the pharyngeal pouch, and a conical papilla pointing caudally and some-
what toward the ventral side. The bulb consists of a loose arrangement
of muscle fibers between which there are very numerous gland ducts con-
taining a fine-grained faintly eosinophilic secretion. These ducts
originate from cell bodies lying in the surrounding mesenchyme and open
within the bulb into a rather small, usually antero posteriorly compressed
cavity, the seminal vesicle (vs). Each vas deferens (vd), which retains
its expanded shape as spermiducal vesicle, enters the penis bulb laterally,
forming a few convolutions within the bulb, and finally narrowing to a
short canal which opens into the seminal vesicle. At the border be-
tween the penis bulb and the penis papilla the seminal vesicle connects
with the ejaculatory duct (de). This is, in its main portion, a highly con-
voluted tube of about even diameter. Only its terminal part gradually
narrows and straightens out in the axis of the papilla, to open to the out-
side at its tip. This opening is encircled by a small collarlike projection of
the papilla. The space between the outer wall of the papilla and the

The genus Planaria 509

convolutions of the ejaculatory duct contains a very loose parenchyma,
often giving the impression of an empty space. This condition, as well as
the fact that the ejaculatory duct does not have the typical shape of a
straight tube makes it possible to speculate whether the ejaculatory duct
is not capable of evagination (like the cirrus of trematodes and cestodes,
or the proboscis of nemerteans). Evidence of a partial eversion was
seen in at least one of the eight specimens sectioned.

The epithelium lining the seminal vesicle consists of columnar to cubi-
cal cells perforated by the numerous gland ducts which have penetrated
the penis bulb from the outer mesenchyme. The lining of the ejaculatory
duct is a cubical, the outer covering of the penis papilla a flattened
epithelium. Both epithelia have associated muscular layers: circular and
longitudinal fibers on the papilla and chiefly longitudinal fibers on the
ejaculatory duct.

The copulatory bursa (b) is a more or less rounded sac situated in the
mesenchyme immediately behind the pharyngeal pouch, lined with a
rather tall glandular epithelium. Its duct or stalk (bd) proceeds from its
dorso caudal side posteriorly as a rather narrow canal, then gradually
widens forming some convolutions above the atrium, and, after narrowing
slightly, bends ventrally and opens into the atrial complex close to the
gonopore. There is, therefore, no enlarged terminal section or vagina
present. The duct is lined with a cubical epithelium which bears cilia at
least in the distal (posterior) part of the canal. It is surrounded by a
well-developed muscle coat of circular fibers adjoining the epithelium,
followed by a layer of longitudinal fibers.

The two ovovitelline ducts, which accompany the ventral nerve cords
in the anterior part of the body, deviate from their course at the level of
the penis, ascend dorsally along the wall of the male atrium and unite in
the space between atrium and bursa stalk. The common oviduct (od) thus
formed runs postero ventrally along the roof of the atrium and opens,
without further differentiations, into the posterior portion of the atrium.
The paired oviducts from the place where they are separated from the
nerve cords, and the entire unpaired or common oviduct receive many
gland ducts with an intensively eosinophilic secretion from the surround-
ing mesenchyme. These glands are generally termed “shell glands”
although their function is doubtful and probably has nothing to do with
the formation of the shell of the cocoon.

The adenodactyl (ad) is a very distinct ellipsoidal or almost spherical
hollow organ situated near the ventral side a short distance posterior to
the gonopore. It consists mainly of a highly muscular covering enclosing
a round cavity. The muscle fibers of the organ are very densely arranged,
mainly in a circular direction, with their cell bodies and nuclei forming
a distinct peripheral layer (Fig. 10, nu). Gland ducts seem to penetrate
the muscular coat from the outer mesenchyme, but do not show up clearly
after staining with hematoxylin and eosin. The lumen of the adenodactyl
opens into a small compartment of the genital atrium which extends pos-
teriorly from the vicinity of the gonopore. There may be a small papilla

506 Proceedings of the Biological Society of Washington

protruding from the adenodactyl into this compartment, or this papilla
may be entirely absent, depending on the state of contraction of the
copulatory complex. The cavity of the adenodactyl is lined with a cubi-
cal ciliated epithelium. In the sections examined, the cavity was usually
empty, without an accumulation of secretions.

Distribution: Planaria occulta has so far been found in only one local-
ity, a hand-dug well, about 4 m deep, just east of the town of Duffield,
Scott County, Virginia, on the property of Mr. Corbett Brown. The
first specimens brought to my attention were collected by Dr. John R.
Holsinger and Mr. Sam Pinkerton on 11 March 1967. They were pre-
served in formalin and showed the reproductive system well developed.
A second lot of sexually mature specimens, collected 8 April 1967 by Dr.
Holsinger, were received in the living state. On 26 November 1968 I
visited the locality, submerged some liver bait in the well, and collected
next morning about 100 specimens of various sizes, the majority having
developed reproductive structures. The water temperature at that time
was 11.8°C.

Ecology: It is difficult to decide whether the hypogean occurrence
of the species is obligate, as the surface waters of the geographic area
have not been examined systematically. The fact that sexually mature
animals were collected in March, April, and November makes it probable
that their sexual maturity is not of a seasonal nature. In the laboratory
the aniamls kept very well in spring water cultures at 14°C and accepted
beef liver and Tubifex worms as food. However, no egg capsules were
deposited during five months of culturing, nor was there any evidence of
asexual reproduction by fission.

Taxonomic position: The general arrangement of the various parts
of the copulatory apparatus and the presence of a hollow adenodactyl
place the species in the genus Planaria. While the remaining species of
this genus P. torva, P. kempi, and P. dactyligera, form a closely related
group with many characteristics in common, the new species occupies a
somewhat isolated position. Apart from the lack of body pigment (which
is also occasionally seen in P. torva, see Reisinger [1963: 685], and in the
cavernicolous subspecies P. torva stygia Kenk [1936: 7]), some features
of the copulatory apparatus deviate considerably from the conditions seen
in the type-species of the genus. Among these, the most outstanding
difference concerns the anatomy of the penis: the presence of a sinuous,
probably eversible ejaculatory duct. The adenodactyl likewise differs in
some details from that of P. torva.

Parasites: All specimens of Planaria occulta sectioned or investigated
in squash preparations were heavily infested with the holotrichous
ciliate Sieboldiellina planariarum (Siebold), a parasite commonly found
in P. torva (see Meixner, 1928: 604, etc.) and other freshwater triclads.
The ciliates were always found in the rami and branches of the intestine,
less often in the pharyngeal pouch, and occasionally in the copulatory
bursa. This latter observation supports the interpretation maintained
chiefly by Steinbéck (1966: 167, etc.) that the bursa is a derivative of

The genus Planaria 507

the intestine which has retained many of the functions of that organ as
well as part of its chemical environment.

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ee gas

Ou

he Ms ; J =
Vol. 82, pp. 559-762 ae X Mig 17 November 1969
De :

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PAPERS PRESENTED AT A SYMPOSIUM ON

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PAST ¢ PRESENT e« FUTURE

DaniEL M. Couen, Special Editor
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Vol. 82, pp. 559-762 17 November 1969

PROCEEDINGS

OF THE

BIOLOGICAL SOCIETY OF WASHINGTON

PAPERS PRESENTED AT A SYMPOSIUM ON

NATURAL HISTORY COLLECTIONS
PAST ¢ PRESENT e FUTURE

DaniEL M. Couen, Special Editor
AND

Rocer F. Cressey, Editor

-

:

TABLE OF CONTENTS
ina tr CUT C tO TN eee se Se ee ee 560

Art and Science as Influences on the Early Development of Natural
History Collections—Phillip C. Ritterbush 561

Vertebrate Fossil Collections—A Fragmentary Document—Nicholas
NEOGEO TU LAL eae es a ere re ee ee _ 579

Fossilk—The How and Why of Collecting and Storing—Ellis L.
Vhoelivelorih ee ee 585

The Role of the National Parasite Collection in Veterinary Parasitology

—Willard W. Becklund 603
The National Collections as Biological Standards—Richard Cowan _— 611
Does Anthropology Need Museums?—William C. Sturtevant _.— . 619

The Role of Museum Collections in Ornithological Research—Richard
STOLL Sa eae a el ra a Se DC a 651

Malacological Collections—Development and Management—Joseph

Rosewater... So OSERA Set OPAIE TS PON RE eS =O ES oes = 665
Automation in Museum Collections—Raymond B. Manning — 671
The Herbarium: Past, Present, and Future—Stanwyn G. Shetler 687

Summary—Daniel Cohen and Ernest Lachner 759

INTRODUCTION

The Biological Society of Washington has periodically as-
sembled to hear scientific lectures and papers since its founding
in December, 1880. The two main functions of The Society
through the years have been sponsoring meetings and publish-
ing a scientific journal, The Proceedings. The membership of
The Society is chiefly composed of systematists, and this has
been reflected in the subject matter of The Proceedings and the
meetings.

As demands for research and service in systematics grow ever
more insistent, collections—the systematist’s singlemost impor-
tant tool—grow larger and their efficient maintenance becomes
increasingly complex and costly. It was considered timely,
therefore, to identify and discuss aspects of the plethora of
problems besetting the managers and users of natural history
collections. To this end The Biological Society devoted its
Autumn 1968 meeting to the topic Natural History Collections,
Past—Present—Future.

The Biological Society is grateful to the speakers, many of
whom are not members, for their conscientious efforts in pre-
paring and presenting a stimulating program. We also thank
The Smithsonian Institution for making available to The So-
ciety facilities for the meeting.

The following papers were presented at the meeting, chaired
by Ernest A. Lachner and arranged by Daniel M. Cohen and
Stanwyn Shetler, which took place on October 11, 1968.1

1 An additional paper not included here entitled “Entomological Collections—The
Dilemma of Success’? was presented by Donald Duckworth, Smithsonian Institution.

Proc, Brot. Soc. Wasu., Vou. 82, 1969 (560 )

ART AND SCIENCE AS INFLUENCES ON
THE EARLY DEVELOPMENT OF NATURAL
HISTORY COLLECTIONS

By Puiie C. RirrersusH
Smithsonian Institution, Washington, D.C.

Cabinets of curiosities and treasure chambers, those early
antecedents of the natural history collection, may seem to us
to have been not at all scientific in their organization or scope
and thus to have had little scientific value. Not until the time
of Linnaeus and Lamarck do we find collections being used
to generate classifications, which has until quite recently been
the primary scientific use to which they have been put. But
before the collection could serve this or any other scientific
purpose it had to be acknowledged that the specimens corre-
sponded to the natural world, that they could represent living
entities as they have actually existed. This belief need not
involve us in questions about the reality of classifications (im-
portant though these have been as determinants of the charac-
ter of modern collections) because it bespeaks a much more
basic presupposition, namely that the external world of living
forms was real and thus might be reliably represented by speci-
mens. It was this basic presupposition that the forerunners
of the natural history collection helped to establish.

The manner in which individuals perceive their surroundings
is greatly affected by their social institutions (Berger and
Luckmann, 1966, 19-34 and 121-22). Ecclesiastical insti-
tutions dominated early medieval Christian Europe to the
extent of claiming and exercising the right to determine which
modes of human experience could be designated as real. Ab-
juring direct means of knowing, the Church aspired to ethical
and spiritual accomplishments which could be experienced
only indirectly, through symbolism or ritual. As aids to attain-

47—Proc. Brot. Soc. Wasu., Vou. 82, 1969 (561)

562 Proceedings of the Biological Society of Washington

ing significant spiritual experiences the Church maintained
extensive visual arrays of symbolic figures and designs, of
which some cathedrals were astonishingly well developed ex-
amples, instructing the people and offering them opportuni-
ties for sustained emotional involvement. “In ages for which
religion and poetry were a common possession, the basic
images lived in the conscious mind; men saw their place and
destiny, their worth and guilt, and the process of their exist-
ence, in terms of them” (Farrar, 1949, 13-14). The material
world was significant only as a symbol for a spiritual reality of
vastly more consequence. To the author of the twelfth-
century De Bestiis a dove had two wings as the Christian had
two ways of life, active and contemplative. Its eyes were
golden because that is the color of ripe fruit and thus of the
wise maturity of the church. Its feet were red for the church
moved through the world with her feet in the blood of martyrs.
Its blue wings reflected thoughts of heaven (Male, 1913, 30).
One of the most widely known works on the significance of
natural objects was the Physiologus, a very ancient bestiary
presenting symbolic interpretations of animal fables. Symbol-
ism gave a rigorous and all-embracing conception of the world
(Huizinga, 1924, 204-5) within which descriptions of natural
entities for their own sake were usually mere “interpolations”
(Crombie, 1952, 8). The naturalistic techniques of illustration
developed during classical times had been virtually lost
(Evans, 1933).

The most prominent works of art in churches throughout
the Middle Ages reflected the symbolic program, but in lesser
works such as decorative architectural details and borders of
illuminated manuscripts the artists of the time were free to
pursue a more independent course. From sources such as the
capitals of columns (Jalabert, 1932) and ornamented books of
hours it appears that there gradually developed during the
thirteenth century a reinvigorated naturalism, reflecting an in-
creasingly widespread ability to perceive the natural object as
an entity in its own right. Around the beginning of the thir-
teenth century the ornamental foliage of capitals of columns
in French cathedrals ceased to be generalized and abstract and
came to portray recognizable species of plants. By comparison

Natural history collection symposium 563

Fic. 1. Anonymous woodcut, “Natiirliche Contrafaytung des Herrn
oder Kiinigs der Chavalette,” signature and date 1542 added in ink. Bor-
der dimensions 18 & 12.5 cm. MS.F13,f88a, Sammlung Wickiana, Zen-
tralbibliothek, Zurich.

to the stylized illustrations of writings on medical topics artists
of the time were equally far advanced in their portrayals of
skeletons and anatomical features. In general these artistic
manifestations of naturalism took place a century or more in
advance of naturalistic descriptions or portrayals of organisms
by learned writers. It would seem to be a consequence of
ecclesiastical control of the most socially important processes
of perception that naturalistic portrayal began as a minority
tendency on the part of artists rather than writers and other
systematic thinkers with whom ecclesiastical authorities were
more concerned. Lynn White, Jr. in an important article
postulated that these artistic developments were the begin-
nings of a later and more general shift in attitudes favoring
naturalism and more concrete representation even of divine
phenomena, as in the eucharistic cult with its tangible sacra-
ments which became prominent at the same time. Such de-
velopments, of course, greatly favored the establishment of
scientific attitudes (White, 1947, 427-31).

A striking example of the distortions of perception induced
by the symbolic view of reality may be found in accounts of
periodic European infestations of the migratory locust with
illustrations portraying it as a demonic and malevolent crea-

564 Proceedings of the Biological Society of Washington

Fic. 2. Monogrammist HW, “Natuerliche Contrafeyhung des gewalti-
gen flugs der Heuschrecken ... ,” dated 1556. Border measurement 18
x 16 cm. MS.F13,f80, Sammlung Wickiana, Zentralbibliothek, Ziirich.

ture. There is an allegorical drawing by Albrecht Durer
(1471-1528) in the Museum at Rennes in which locusts are
depicted as devils writing script and carrying various sinister
objects (Blanck, 1957, 6). Such an illustration shows the in-
fluence of prevailing theological conceptions of the locust as
an instrument of divine vengeance. During the plague of 1542
one observer claimed to find the words IRA DEI on the wings
of locusts, which he took as evidence that they were indeed
messengers of divine wrath (Schoénwialder, 1960, 413). After
the infestations of 1542 and 1556, each extending through wide
areas of Italy and central Europe, woodcuts were made show-
ing locusts as fabulous beings with exaggerated antennae,
webbed feet, a forward-pointing spiral appendage (in the 1542
drawing ), and brush-like tails (Fig. 1 and 2). These illustra-

Natural history collection symposium 565

Fic. 3. Realistic depiction of migratory locust by Pisanello, ca. 1430.
Musée de Louvre, Paris. Photo credit: Cliché des Musées Nationaux.

tions were published as parts of broadsides printed to carry
news of the locust plagues and thus may be taken to represent
attempts to record the events. There is strong confirmatory
evidence that there indeed were plagues of locusts when re-
ported (Baccetti, 1954, 278; Waloff, 1940, 225) yet visualiza-
tions strayed exceedingly far from their objective basis. As late as
the middle of the sixteenth century it was possible for a would-
be chronicler to have before him a locust yet perceive and
record a chimera, as the socially derived mode of perception
imposed itself upon the data of experience. The early drawings
of the locust were frequently so schematized as to be unrecog-
nizable. One of the most experienced students of medieval il-
luminated manuscripts reproduces two drawings from the late
thirteenth and early fourteenth centuries in which peasants are
filling sacks with migratory locusts. The captions are errone-
ously given as “Man and butterfly, pursuing with hood.” (Ran-
dall, 1966, PI. LXXI, figs. 342, 343).

In the Louvre there is a drawing executed over a century
earlier, by Pisanello (1380-1456), in careful naturalistic detail,
clearly recognizable as Locusta migratoria and lacking any of
the fantastic features attributed by the artists of the later wood-

566 Proceedings of the Biological Society of Washington

cuts (Fig. 3). The naturalistic illustration had been far ad-
vanced for its day and the later woodcuts may be taken to
show a persistence in popular culture of the fabulous tenden-
cies in depictions of creatures influenced by prevalent medieval
concepts of reality. This interpretation posits a gradual change
in modes of perception by which naturalism appeared first as
an esthetic motive in the decorative arts and then grew in im-
portance until it became the basis for more accurate scientific
representations of creatures based upon direct observation un-
hindered by conceptual distortions.

Leonardo da Vinci (1452-1519) exulted in the knowledge he
gained from direct observation. It is significant that the most
profound Renaissance conception of the scientific value of
naturalistic perception was that of an artist, who indeed con-
ceived of painting as the highest form of knowing. Leonardo’s
avowal that “All our knowledge originates in our senses”
(Stites, 1968, 222) sharply contrasts with the verbal procedures
by which contemporary academicians still sought to substan-
tiate their beliefs. Leonardo praised the power of drawings to
describe a “whole arrangement,” far superior to verbal de-
scriptions which conveyed “but little perception of the true
shapes of things” (Zubov, 1962, 57). From 1485 he had con-
ducted serious anatomical studies based upon numerous dis-
sections. He advocated consecutive drawings to show how
different systems composed an organ and also sequential
drawings to depict the same structure from several directions,
and he tried also to represent living things in their dynamic
aspect. His ideal was the geographic atlas showing all major
provinces of a subject. The artist must progress beyond naive
perception to discerning visual examination of objects. He
must “know how to see” (saper vedere). Leonardo was
especially contemptuous of beliefs that immaterial spirits, lack-
ing extension and the capacity to exercise force, could inter-
vene in the everyday world. His observations clearly demon-
strate the important consequences for scientific knowledge
which would follow from learning to see.

One may perhaps mark the turning point in the application
of naturalistic perception to biology in the work of Vesalius
(1514-1564 ). In the well-known scene of an anatomical theater

Natural history collection symposium 567

that appears as the title page of De humani corporis fabrica
(1543) there is a bearded man holding a closed book while
pointing to the dissection in progress as though to admonish
a nearby student that more is to be learned from reality than
books. Indeed, it required only the most cursory observation
to demonstrate that men do not lack a rib even though Moses
wrote that God took one from Adam or that the human liver
does not have the five lobes which Galen ascribed to it.

We might note that the most important forerunner of Vesa-
lius, Giacomo Berengario da Carpi (c. 1460-1530), was praised
by Cellini for his interest in art and possessed a considerable
art collection. The splendid woodcuts commissioned and per-
haps partly executed by Vesalius established the importance
of biological illustration, and they reveal something of their
artistic legacy in the landscapes of the Euganean Hills near
Padua drawn in the background of the plates of “muscle-men,”
as well as in the poses of the figures, taken from antique
statuary. Perhaps mindful of the dissections carried out by
the artists Antonio Pollaiuolo (1429-1498) and Benozzo Goz-
zoli (1420-c. 1497), as well as Leonardo, the recent biographer
of Vesalius observes that “The impulse to naturalistic anatom-
ical depiction seems to have come from the art world rather
than the medical.” We should also note his observation on
the extent to which Vesalius owed his success to the reviving
naturalistic mode of vision: “Vesalius had an extraordinarily
well-developed visual sense, and it is apparent in his verbal
descriptions of anatomical structures” (O'Malley, 1964, 18
and 118). The Historia animalium of Conrad Gesner (1551)
and De historia stirpium of Leonhart Fuchs (1542), both pro-
fusely illustrated works, were published at about the same
time, indicating that the use of realistic illustrations had be-
come established (Nissen, 1963; Ziswiler, 1965; Blunt and
Stearn, 1950).

The ability to discern and portray accurately the charac-
teristics of the form of organisms, a talent at odds with the
prevailing official mode of the time, owed its origin to artists
and illustrators. The further extension of this ability in so-
ciety would depend upon the extent to which men could learn
to see in naturalistic rather than in symbolic terms. The phenom-

568 Proceedings of the Biological Society of Washington

enological foundations of biological science were laid by
naturalistic artists several centuries before the prevailing views
came to ascribe the force of evidence to direct observation and
objective portrayal of specimens from nature. Thus we should
be on the lookout for new institutions serving to apply the
artists’ mode of perception to the social enterprise of ascribing
reality to man’s experience. The cabinet of curiosities, the early
forerunner of the natural history collection, served a mediating
function of this kind.

In the evolution of natural history collections the visual
arts played a role which seems to have been central but which
is difficult to define. There were no public museums until the
eighteenth century. Scientific collections evolved slowly from
the private treasure chambers of robles and kings. Virtually
the only natural objects found in these collections were fabu-
lous or prized for their rarity. In the collection of Jean, Duc
de Berry (1340-1416) there was a wonder cabinet with giants’
bones, sea monsters, carved crystals, and some genuine articles
such as ostrich eggs and polar bear skins. By the sixteenth cen-
tury there were about a dozen outstanding large collections of
princely treasure such as that of Archduke Ferdinand of Tirol
(1520-1595) at Schloss Ambras (Schlosser, 1908). In these
collections natural history objects were combined with gems
cut into natural forms, montages of shells, and decorative items
made from natural substances. The word cabinet is used some-
times of the collections as a whole and sometimes of the chests
containing smaller items. The Kunst-und-Naturalienkammer
set up by the Elector Augustus I (1530-86) of Saxony com-
prised seven rooms of the Royal Palace in Dresden, with works
of both fine and decorative arts intermingled with natural his-
tory objects (Wittlin, 1949; Schuster, 1929; Murray, 1904;
Bedini, 1965).

One of the most elaborate of the cabinets ever built to store
such intermingled collections of nature and art objects is pre-
served in Uppsala. It was made by Philip Hainhofer of Augs-
burg (b. 1578), whose paintings and collages are occasionally
remembered as examples of optical illusions, many based upon
natural form. He was a dealer in natural rarities and art who
oversaw the preparation of one celebrated cabinet in 1617

Natural history collection symposium 569

Fic. 4. Gem and art peak of the Gustavus Adolphus Kunstschrank
(1625-26). The vessel is 42 cm long and the work of H. C. Lencker, an
Augsburg silversmith. From Béttiger, 1910, Plate 12.

for Duke Philipp II of Pomerania, which was brought to Ber-
lin to hold part of the royal collection and destroyed during
World War II (Lessing and Briining, 1905). The Uppsala
cabinet, which was prepared in 1625-26, rises in several tiers
of ebony drawers and contains numerous doors opening onto
facades of cameos and rare woods. It is crowned by a carved
coconut, coral, and silver drinking vessel with statuettes of
Neptune and Venus atop a distinctive montage of minerals
(quartz, citrine, hematite, barite, ores, and semiprecious
stones ) and shells ( Fig. 4).

In the centuries following wealthy private collectors and

570. Proceedings of the Biological Society of Washington

Fic. 5. Works of art and natural objects combined in a seventeenth-
century collection, painted by Frans Francken the younger (1581-1642),
“Eine Kunst und Rarititenkammer” (undated), 74 * 78 cm, Kunsthis-
torisches Museum, Vienna.

scholars also formed collections. Here, too, we find coins and
other antiquities, shells and marine specimens, gems, and
paintings indiscriminately jumbled together, as in the remark-
able painting by Frans Francken the younger (1581-1642)
showing a gentleman’s collection and its owner discoursing
over books with his friends in an adjoining room (Fig. 5). In
the collection of Ulisse Aldrovandi (1527-1605) at Bologna
works of art were arranged as ethnological curiosities or as
examples of the materials of which they were made while
natural objects and imitations were placed together (Schlosser,
1908, 108). An illustration of the collection of the pioneer
marine biologist Ferrante Imperato (1550-1625) in Naples
shows one wall lined with cabinets for works of art and the

Natural history collection symposium 571

Fic. 6. Objects of art and nature combined in an early collection. Frontis-
piece, Ferrante Imperato, Dell’historia naturale . . . (Naples: C. Vitale,
1599).

ceiling covered with marine productions arranged without re-
gard for their biological affinities (Fig. 6). In 1725 the collec-
tion of Sir Hans Sloane (1660-1753), which was to form the
nucleus of the British Museum, included 5497 minerals and
fossil substances, 804 corals, 8226 vegetable substances, 200
volumes of dried plants, 3824 insects, 3753 shells, 1939 echi-
noids, fishes, crustaceans, etc., 568 birds and 185 eggs, 1194
quadrupeds, 345 reptiles, 507 human objects, 1169 miscella-
neous artificial and natural objects, 302 antiquities, 81 large
stone seals, 319 pictures, 54 mathematical instruments, 441
vessels and carved mineral objects, 136 illuminated books,
20,228 coins and medals, 580 volumes of prints, and 2666 manu-
script volumes (Murray, 1904, I, 137-38).

The inclusion of the fine and decorative arts in these collec-
tions affords a clue to the intricate cultural change that was
occurring. The princely collection with fabulous or exceed-
ingly rare animals was gradually succeeded by a collection
representative of the animal or plant kingdom. The decorative

572 Proceedings of the Biological Society of Washington

objects so important to the early collections dwindle by propor-
tion until by the eighteenth century one finds collections made
up exclusively of natural objects. It would seem that the works
of art in the collections functioned as catalysts in an uncon-
scious transfer of authority from the artists’ perception to the
naturalists’ reliance upon the objects themselves. We have to-
day none of the collections as they were; objects of art and
nature once regarded together have become the separate re-
sponsibilities of distinct departments in modern museums
(Hutchinson, 1965.) Further study of inventories and de-
scriptions of sixteenth and seventeenth-century collections is
surely desirable to clarify and define the effect of art works
upon the perception of natural objects and changing concep-
tions of reality as they have represented it. Such a correla-
tion of the contents of collections with the conceptual develop-
ment of biology would be a welcome contribution to the
history of scientific thought.

Toward the end of the sixteenth and throughout the seven-
teenth century realistic still-life paintings of flowers and insects
became immensely popular in the Low Countries ( Bergstrém,
1956; Wamer, 1928; Bernt, 1948). Paintings by Jan Brueghel
the elder (1565-1625), Ambrosius Bosschaert the elder (1573-
1621), Roelandt Savery (1576-1639), Daniel Seghers (1590-
1661), Jan Davidsz. de Heem (1606-1653), Otto Marseus van
Schrieck (c. 1619-1678), Abraham Begeyn (c. 1637-1697),
Abraham Mignon (1640-1679), Rachel Ruysch (1664-1750),
Jan van Huysum (1682-1749), and others frequently portrayed
flowers in precise detail with recognizable species of insects
situated near them in lifelike poses, while snails and snakes
often appear. One of the earliest and most interesting of these
painters was Georg Hoefnagel (1542-1600), whose works
showed many exotic insects brought to Europe for the first
time (Kris, 1927; Bergstrom, 1963).

The style of these works is usually termed “scientific natu-
ralism.” One leading scholar has attributed the realism of Dutch
and Flemish flower painting to the “philosophy which claimed
that the quality of reality belongs exclusively to the particular
things directly perceived by the senses” (Panofsky, 1953, I, 8).
The flowers frequently symbolize mortality and sometimes the

Natural history collection symposium eee)

Fic. 7. The painted surface of the Smithsonian cabinet of curiosities,
attributed to Jan van Kessel. 42% in * 26% in. External marquetry dec-

oration appears above and below the painted surface.

insects are allegorical representations (Bergstrom, 1955), but
the overwhelming impression created by these numerous works
is one of fascination with their immediate colorful subject
matter. They thoroughly document the force and persistence
of naturalism as an artistic motive throughout the period of
the development of the natural history collection.

In 1964 the Smithsonian Institution acquired some months
after its sale at Sotheby’s (March 11, lot 88) an exceptionally
interesting work in this genre which serves to remind us of the
close links between naturalism and cabinets of curiosities ( Fig.
7). It is a seventeenth-century veneer and marquetry cabinet,
an unsigned work of Flemish or English craftsmanship, with
ten drawers and a central door panel whose veneer surfaces are
painted white and on which appear scores of insects, painted
approximately life-size, after the manner of the well-known
Flemish still-life painter Jan van Kessel (1625-1679), who is
well represented in major European museums. Many of the

574 Proceedings of the Biological Society of Washington

individual insects and even their arrangement in the panel
compositions are identical to those in signed works by van Kes-
sel. The entire composition closely resembles that of a set of
seventeen paintings on copper signed and dated 1658 bought
by the Amsterdam firm Gebr. Douwes in England in 1923 and
sold by them to a Mr. van Valkenberg in 1924. This set is
probably the same as that sold by the Fievez firm in Brussels
in 1935 and that exhibited by the Hallsborough Gallery in Lon-
don in 1956, and since sold to an anonymous buyer (personal
communications from Evert J. M. Douwes and the Hallsbor-
ough Gallery; also Hallsborough, 1966). A separate, virtually
identical set was exhibited in Amsterdam in 1934 by the firm
of P. de Boer and then broken up (personal communication,
P. de Boer). Both sets on copper were probably prepared for
the fronts of cabinets, either as decoration or explicit commen-
tary on cabinets of curiosities.

The Smithsonian cabinet is not as intricate in detail as most
van Kessels; it was probably copied in England from one of the
sets on copper or possibly executed in van Kessel’s own studio
in Antwerp. The latter would be more likely if the place of
the cabinet’s manufacture could be established as Flanders,
but its manner of decoration was virtually an international
style, so that it is very difficult to assign individual pieces to
one country or another. The dimensions of the Smithsonian
cabinet are more regular in inches than in pieds and pouces, the
system of measurement in use on the Continent at the time,
which suggests that it was fabricated in England. At any rate,
its design clearly reflects van Kessel’s work of 1658 and the
tradition by which naturalism had come to be associated with
cabinets of curiosities.

The insects and plants, as was true of most work of the genre,
were almost certainly copied from sketchbooks (a practice that
enabled artists to produce their works throughout the year, not
just when flowers were in bloom and insects on the wing).
It is also of interest that van Kessel executed works in which
creatures were portrayed almost as in the dioramas of museums
(usually considered a nineteenth-century innovation). In the
Museé de Dijon are two undated works of this type: one,
“L’eau,” shows a seal, giant squid, and numerous fish on a

Natural history collection symposium bio

Fic. 8. The central panel of the Smithsonian cabinet of curiosities,
attributed to Jan van Kessel. 12° in * 15% in. The figure derived from
the locust woodcuts appears in the lower center.

beach; the other, “La terre,” shows stags, peacocks, roses and
other plants, and two hawks tearing at a dead game bird. An-
other painting sold by the firm of Nystaad in Lochem in 1947,
entitled “The night,” portrays a lively group of bats, badgers,
and wildcats in a nighttime landscape.

It is in the central panel of the Smithsonian cabinet, with

576 Proceedings of the Biological Society of Washington

exotic insects and arachnids from the Americas (Fig. 8) that we
encounter an image which reminds us of the progress made
toward naturalistic representation in the century or so preceding.
Here, slightly altered, but with unmistakable thickened an-
tennae and forward-pointing spiral appendage is the migratory
locust figure seen previously in the sixteenth-century broad-
sides! Before seeing those earlier illustrations in the Ztirich
library I had supposed that this was an illustration of a “hum-
bug” fabricated by curio merchants (Misson, 1699, I, 134-35;
Ripley, 1965; Ritterbush, 1964, 145 n.). To find the sixteenth-
century illustrations of the locust was to discover an un-
expected element of continuity linking the fabulous images
of a symbolic age to the progress of realistic natural knowledge
based upon the objects themselves, but only as a single sur-
vival amidst an array of realistic portrayals. If the collection
was gradually transformed from an artistic aggregation to a
purposeful instrument of scientific inquiry it was because men
of science had learned to see, largely as a result of the vivid ac-
complishments of artists who had so far preceded them in em-
ploying naturalistic vision. A treatise on museums and collec-
tions published in the early eighteenth century included as its
dedicatory legend a verse which seems aptly to summarize this

history (C. F. Neickelius, 1727):

What in this world can more delight

Than the nobility of creatures studied as they really are?
What can excite joy and wonder in the soul

More than viewing the reality of nature?

LITERATURE CITED

Baccetti, Baccio. 1954. Storia delle infestazioni di ortotteri in Italia
con particolare riguardo a quelle verificatesi in Toscana.
Redia. Sr 2, 39: 275-289.

Bepint, Sitvio A. 1965. The evolution of science museums. Technology
and Culture, 6: 1-29.

BerGER, PETER L., AND THOMAS LUCKMANN. 1966. The social construc-
tion of reality. Doubleday, New York. Anchor Books, 1967.

BercstroM, INcvArR. 1955. Disguised symbolism in ‘madonna’ pictures
and still life. Burlington Magazine. 97: 303-308 and 342-
AQ,
1956. Dutch still-life painting in the seventeenth century.

Natural history collection symposium 577

Trans. Christina Hedstrom and Gerald Taylor. Thomas Yosel-
off, New York.

1963. Georg Hoefnagel, le dernier des grands miniaturistes
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VERTEBRATE FOSSIL COLLECTIONS—
A FRAGMENTARY DOCUMENT

By Nicnoxias Horron III
Smithsonian Institution, Washington, D. C.

The fossil record plays a unique role in the study of biology,
for it provides our only appreciable access to the time dimen-
sion of evolution as an historical process. To expatiate mo-
mentarily upon the obvious, the most nearly universal charac-
teristic of the fossil record is the fact that it is fragmentary.
The causes of this fragmentary nature—structure and mode of
lite of plant or animal, age, and pure chance—also introduce a
bias into the fossil record. We need not concern ourselves with
the causes, but the fact that the record is fragmentary and bi-
ased has a strong influence on the study of paleontology, par-
ticularly of the vertebrates, and on the role of museum collec-
tions in this study.

In practice there are two vertebrate paleontologies, one con-
cerned with animals which have lived since the end of the
Mesozoic Era, and the other with animals that became extinct
before that time. The difference between the two is deter-
mined by three factors, one biological, the degree of similarity
of the organisms to living forms, and a second geological, the
degree of similarity of past to present physical circumstances of
the earth’s surface. The third factor, completeness of the rec-
ord, is a product of both biological and geological influences.

The Cenozoic Era, the approximately 70 million years that
have elapsed since the end of the Mesozoic, is often called the
Age of Mammals, in reference to the fact that the dominant ter-
restrial vertebrates of this interval are mammals very similar in
general to living forms. Lineages of the major orders of living
mammals can be traced with a high degree of confidence in the

48—Proc. Brot. Soc. Wasu., Vou. 82, 1969 (579)

580 Proceedings of the Biological Society of Washington

changing faunas of the Cenozoic. The younger the faunas in
question, the more directly can their components be compared
with living animals, and although this comparison becomes
somewhat more difficult as one goes back in time, the mam-
mals of even the earliest Cenozoic are sufficiently similar to
those of the present day to afford a basis for direct comparison.
This is also true of Cenozoic amphibians (frogs and salaman-
ders ) and reptiles (crocodilians, lizards and snakes, turtles, and
scattered relatives of Sphenodon).

Terrestrial vertebrate faunas prior to the Cenozoic were
dominated by reptiles, most of the Mesozoic by dinosaurs, and
the late Paleozoic and earliest Mesozoic by synapsid, or mam-
mallike reptiles. Each of these groups was preeminent for about
130 million years, almost twice as long as the mammals have
thus far enjoyed their supremacy. The most striking characteris-
tic of these animals is the difference between them and the
reptiles—or anything else—living today. Dinosaurs are often
compared with birds, to which they are closely related, and
synapsids can be compared with their descendants the mam-
mals, or with unrelated reptiles such as turtles, with which they
have many habitus features in common. But this is a far cry
from comparing an early Cenozoic horse with Equus, or a Mio-
cene arctoid carnivore with living dogs or bears. No terrestrial
tetrapod of the present is closely comparable to either di-
nosaurs or synapsids in its general organization, in the way it
makes its living. As a consequence, one is restricted to methods
of classic comparative anatomy in working out relationships of
pre-Cenozoic tetrapods, and resolution of all problems, whether
taxonomic or functional, must be based for the most part upon
the remains themselves, with only peripheral or analogical ref-
erence to living animals.

Geological aspects of Cenozoic time are similarly much
more nearly comparable to present-day conditions than are
those of earlier time. The major subdivisions of the fossil rec-
ord, the Paleozoic, Mesozoic, and Cenozoic Eras, are related in
some degree to long-term phases of mountain-building (tec-
tonic cycles) over large parts of the earth. The cycle in which
we find ourselves today was initiated at the beginning of the

Natural history collection symposium 581

Cenozoic and is still active. It has molded and continues to
mold the general configuration of land masses (including major
topography and drainage ), and in doing so controls deposition
of sediments and preservation of fossils.

Presumably the tectonic cycles of the Paleozoic and Mes-
ozoic exercised the same influence over physical conditions on
continental surfaces and over preservation of the faunas of
those times. But the earth movements of each tectonic cycle
result in the destruction of large parts of the features formed
during preceding cycles, and in consequence much of the Pa-
leozoic and Mesozoic record has been lost. Because present-
day tectonics are essentially a continuation of the Cenozoic
cycle, a far larger proportion of the Cenozoic terrestrial fauna
is still preserved and exposed on the surfaces of all continents
except Antarctica and perhaps Australia. Cenozoic faunas
therefore tend in general to be more nearly complete and con-
tinuous than those of earlier time.

Destruction is selective. The higher the land, the more
quickly it is eroded, and upland faunas are therefore rare even
in the Cenozoic, except in its most recent phases. With a few
notable exceptions, upland faunas are unknown in the Paleo-
zoic and Mesozoic.

Tectonic activity of the current cycle has broken up the rec-
ord of earlier eras both temporally and geographically. Except
for bits and pieces, the long history of the dinosaurs is adduced
from three segments of time totalling a good deal less than half
their overall record. The earliest segment is that of the Upper
Triassic, of perhaps 5 million years’ duration, best represented
in South Africa, Brazil and Argentina, western United States,
and western Europe. The second segment, straddling the bound-
ary between Jurassic and Cretaceous, lasted no more than 15
million years and perhaps as little as 5 million, and is best rep-
resented in western United States, western Europe, and Tan-
zania. The third segment is that of the Upper Cretaceous, of
about 30 million years’ duration, best represented in western
United States and Outer Mongolia.

The history of synapsid reptiles as such (omitting Mesozoic
mammals ) extends essentially from the origin of reptiles some-

582 Proceedings of the Biological Society of Washington

time in the early Pennsylvanian to the end of the Triassic. The
record is perhaps more nearly continuous temporally than that
of the dinosaurs, but is sharply broken geographically. Ap-
proximately the first half, to the end of the Lower Permian, is
best represented in the United States, while the second half is
preserved in Russia, South Africa, Zambia, and Tanzania, and
Brazil and Argentina.

The general effect of the characteristics of Cenozoic tetra-
pods and their record is to permit taxonomy and faunistics of the
organisms to be studied in considerable detail. Species popula-
tions can often be recognized on the basis of preserved mate-
rial, and confirmed, at least by analogy, by comparison with liv-
ing populations. At higher stratigraphic levels, studies of rates
of origin and longevity of genera and species in terms of abso-
lute time are possible. In general, the most significant taxo-
nomic work is concentrated below the level of order. Because of
the relatively continuous record, studies of distribution are
more meaningful, and the question of past migration can be
approached directly. The occasional presence of such paleon-
tological exotica as upland faunas gives students of this time
period a better perspective for explicitly ecological faunal stud-
ies. Around its periphery, vertebrate paleontology of the Cen-
ozoic merges imperceptibly into the more strictly biological dis-
ciplines of mammalogy and herpetology.

Few of these approaches are effective in the study of pre-
Cenozoic tetrapods. Disjunction of the record makes questions
of distribution and migration almost meaningless, for although
we know where the animals were, we can never be sure of
where they weren't. Because dinosaurs and synapsid reptiles
are so different from living animals in morphology and biologi-
cal requirements, and because we cannot be confident that the
natural sampling of fossilization has preserved biologically rel-
evant populations, in most cases we cannot confidently recog-
nize reproductively isolated natural populations in the fossil
material. Species designations are used to keep the picture con-
sistent with neozoological practice, but in general the lowest
operational taxonomic unit appears to correspond most closely

Natural history collection symposium 583

to the genus of neozoology. Much of the significant work is
concentrated at about the level of subclass.

Study of Paleozoic and Mesozoic vertebrates is therefore
broad-brush paleontology. Although its low-level taxonomy is
shaky, it provides an overall view of vertebrate evolution which
since Darwin’s time has gone far to document true relationships
between vertebrate classes. Another approach, which has
become more feasible in recent years as more material has
become available, is the study of the functional anatomy of
these outlandish beasts, which ultimately may provide insight
into the selective forces that produced differentiation to such
a high taxonomic level. Vertebrate paleontology of the Paleo-
zoic and Mesozoic draws most heavily from comparative anat-
omy among the strictly biological disciplines, both in the
classic approach and (by analogy) in studies of function.

The value of museum collections to vertebrate paleontology
of whatever period is directly related to the fragmented qual-
ity of the record, for we can never predict what unprepossess-
ing scrap of a fossil will next fill a gap in our knowledge. For
Cenozoic specialists, identifiable bits often provide valuable
data extending temporal or geographic range of mammal spe-
cies. Fragments of crocodilians, turtles, and lizards identifia-
able no more closely than to subclass may contribute to the
understanding of past climatic conditions, for these reptiles
were presumably more restricted than mammals by climatic re-
quirements. For the student of dinosaurs or synapsids, bits and
pieces of ear, braincase, or jaw have contributed to resolution
of problems of function and of high-level taxonomic relation-
ships. This principle is also valid, of course, with respect to
continuing field programs. One can predict only very generally
what he will find in a given area, and it is only by sustained
methodical collecting that these unexpectedly valuable pieces
of the jigsaw puzzle accumulate.

In spite of its incompleteness, the fossil record is so enormous
that no single institution can hope to cover more than a small
part of it comprehensively, and few institutions are large
enough to have a completely representative collection in all
areas. Economic factors dictate that most museums that include

584 Proceedings of the Biological Society of Washington

vertebrate fossils concentrate on a more or less regional cover-
age. As a consequence, collections themselves represent ad-
ditional fragmentation of available material.

The reason that vertebrate paleontology has been so success-
ful in piecing together the torn-up manuscript with which it
must work is that the material has in fact been available, if
scattered. Great strides have recently been made in the inter-
esting transitional areas between amphibians and reptiles, and
between reptiles and mammals. Although both were triggered
by discovery of new specimens, both were properly consoli-
dated and documented by exhaustive reexamination of old
material, some of it having been available for about 150 years.
These developments have stimulated activity in these and re-
lated areas, and more information may be expected momen-
tarily, but if the potential of this sort of work is to be realized,
collections must remain readily available. The question is not
only how to make room for new and significant material, but
how to do this and at the same time keep existing collections
efficiently accessible.

A final point to emphasize is that for the decipherment of the
morphology and general organization of extinct vertebrates,
paleontologists are restricted to a single organ system, the
skeleton. Fortunately, the vertebrate skeleton is biologically
plastic, and readily reflects the former presence of many soft
parts, as well as certain aspects of growth and development.
But in order to interpret these features effectively, the verte-
brate paleontologist is very dependent upon collections of pre-
served specimens of present-day animals. For some problems,
such as direct comparison of populations, he requires skeletons
or suites of skeletons. For others, such as those involving com-
parative anatomy, he requires alcoholic specimens for detailed
dissection. In summary, then, because of the incompleteness
of primary materials, the continuing effectiveness of vertebrate
paleontology requires that as much material as possible be
available, not only fossils, but also relevant Recent specimens.
In this field it is possible, in large measure, to compensate for
the lack of what we can’t get by accumulating an abundance of
what we can.

ord

L

FOSSILS—THE HOW AND WHY OF COLLECTING
AND STORING

By Exzis L. YOCHELSON?
U.S. Geological Survey, Washington, D.C.

INTRODUCTION

Fossils, like eggplant and okra, are a matter of taste in the
American community of naturalists. They are loved by a tew
specialists, tolerated by a few more broad-minded individuals,
actively disliked by some extremists, but essentially ignored by
the bulk of the populace. Accordingly, it is appropriate to re-
view the ways that fossils arrive at museums, and their result-
ing fate, if only to bring these remains of organisms into the
lifestream of natural-history collections.

No one knows how many fossils are still to be recovered from
sedimentary rocks; no one even knows the far smaller total of
the millions of fossils already collected and scientifically stored.
Accordingly, this lack of knowledge provides an ideal oppor-
tunity for the fabrication of fact. The combined U.S. Geological
Survey-Smithsonian Institution collection housed in the Mu-
seum of Natural History of the Smithsonian at Washington cer-
tainly contains more fossils than any other collection in North
America and probably in the world. It is probably safe to add
that many universities store only a token number of fossils in
collections, though there are some impressive lots on a few se-
lected campuses. Most collections outside Washington, D. C.,
are in State geological survey collections or in a limited num-
ber of major but somewhat smaller museums. Some oil com-
panies maintain large numbers of microfossils, but these are
out of the public domain.

Using a dirty crystal ball, one arrives at the figure of less
than 25 percent and greater than 10 percent for the part of the

1 Publication authorized by the Director, U. S. Geological Survey.

49—Proc. Biot. Soc. WaAsH., Vou. 82, 1969 (585)

586 Proceedings of the Biological Society of Washington

Nation’s fossil collection that is in Washington, D. C. Roughly
20 percent of the Museum of Natural History storage space in
Washington is devoted to fossils of the U. S. Geological Sur-
vey-Smithsonian Institution collection. However, this particular
fossil collection has a far greater significance than just as a
large percentage of the total American scientific material, for
the Washington-stored fossils contain more specimens that
have a documentary function than any other collection. Wash-
ington-based persons may be provincial and still do a fair job
of study, but sooner or later paleontologists from other areas
should visit Washington to look at types and special collections.

It is obvious from the preceding statements that the remarks
expressed in this note are necessarily my own. Without attempt-
ing to degrade the variety of opinion in other fields of natural
history, each reader should be informed quite clearly that pa-
leontologists are highly individualistic in all facets of their ac-
tivity. The reader is hereby warned that future statements
made are entirely unsupported opinion. They apply mainly to
fossils stored in Washington, and to their custodians, but might
be more generally applicable if the underlying biases happen
to strike a local and familiar chord.

Wuy Co.Liecr?

There is little sense in beating the dead horses of inherent
curiosity, pushing back frontiers of science, search for the un-
known, and other cliches to answer the question of why a per-
son collects natural-history objects. A paleontologist collects
fossils because he is professionally interested in them; others
collect fossils to derive information or enjoyment from their
possession. The paleontologist occupies an intermediate posi-
tion between the pure compiler of geologic data and the pure
lover of objects.

The latter might be mentioned first, though he does not
deserve such harsh condemnation as mere “object lover.” Ama-
teur collectors are rare in the United States; the semi-pro who
supplements his income by sale of fossils is even rarer. They
may gather important collections, and they should be en-
couraged, but their overall contribution is negligible, especially

Natural history collection symposium 587

when compared with the contribution of the amateur in Eu-
rope. There are probably fewer qualified amateur collectors
now than in past years; the era between the Civil War and
World War I was their heyday.

Field geologists and stratigraphers form the large group that
uses information derived from study of fossils. The former are
concerned with rocks of varying ages in a limited area; the lat-
ter are concerned with rocks of a more restricted age over a
broader area. Both are concerned with questions of time or
depositional environment of the rocks, and they pick up fossils
to obtain evidence bearing on these points. It is my guess that
more than 50 percent of the fossils in Washington were col-
lected primarily to answer the problem of age of rocks. The
percentage may be only slightly less in other large collections.

Paleontologists suffer many disadvantages in their studies
because their specimens are incomplete in a variety of ways
when compared with the biota that may be obtained in the
Holocene. However, they do have one remarkable advantage
over the neontologists in that collecting fossils is a four-di-
mensional operation involving latitude, longitude, altitude, and,
uniquely, time. Others may write learned tracts on evolution-
ary theory, but only the paleontologist can collect one form at
the bottom of a sequence of rock and another, related but
slightly different, at the top. This element of time is the key
factor in paleontology and is a dimension lacking in neontol-
ogy.

In a crude way, one can draw a parallel between the field
geologist awaiting the word of the paleontologist as to the age
significance of a petrified form, and a quarantine inspector
awaiting the word of an entomologist as to the identity of an
insect before deciding whether to permit entry of a boatload
of bananas. Another sort of time factor also enters here, for
most paleontologists are of the opinion that anyone else’s col-
lection of fossils not only can, but should, wait to be examined.
Paleontologists in museums, surveys, and groves of academe,
often in good conscience, may delay months and even years in
producing an answer to an inquiry about the age of a rock;
bananas cannot wait that long. In contrast, the paleontologist

588 Proceedings of the Biological Society of Washington

employed by the oil company currently drilling a well is under
even more pressure than a banana inspector.

Paleontology is closely tied to geology. In the past, although
vertebrate paleozoology in toto and paleobotany in part were
ignored by the field geologists as sources of useful data, inver-
tebrate paleozoology was bound nearly hand and foot to the ef-
fort of age determinations. For the past few decades, this tie
has loosened as the principal masses of sedimentary rocks were
given relative age dates of moderate precision. This has also
come about because of a shift of interest toward other prob-
lems in geology and a shift toward more biological topics in
paleontology.

Under no circumstances should these remarks be interpreted
to mean that the job of even approximate dating by fossils has
been completed, or that it has been even locally accomplished
with maximum precision. More accurate relative dates remain
a prime job for the paleontologist. This close association of
fossils and stratigraphy has been overemphasized in the past
and is underemphasized in the present. As in many other situa-
tions, the middle ground is probably the route to pursue. Even
with the relaxation of the stratigraphic tie, the paleontologist
still obtains a large fraction of his material from the nonpaleon-
tologist. The relation of geologist to paleontologist is certainly
closer and more mutually meaningful than that of, say, geneti-
cist to entomologist. Such relations should be encouraged.

How To Co.Lecr

When one asks a paleontologist how he collects fossils, the
answer is generally a curt reply such as “meticulously.” There
are a variety of techniques, governed mainly by the kind of fos-
sils and the kind of sediment which encloses them. Some peo-
ple swear by a l-pound hammer with a chisel end and a 14-
inch handle; others swear at it. So many common-sense features
are involved in collecting that a brief general summary on the
subject was reviewed as being “downright inane.”

In spite of this opinion, I believe that much remains to be
discussed and written on the subject of fossil collecting. Al-
though professional collectors have been employed permanently,

Natural history collection symposium 589

this luxury is largely a thing of the past in the United States.
Today, people continue to provide inadequate locality informa-
tion when they submit collections of fossils for examination and
do such silly things as write labels in water-soluble ink. If any-
thing, the ability to obtain useful fossils, ship them, and have
the collection arrive in reasonable shape and containing the
proper information has lessened as interest in fossils has de-
clined among nonpaleontologists.

Collecting may be reduced to two fundamentals. First, find
a specimen, and second, retain it at least for a significant time
interval. Expressions commonly heard are that collections were
made, but after several years of just having them take up space,
the fossils were discarded. Alternatively, one hears of the pro-
verbial mountain slope littered with fossils, but they were not
collected because the age of the formation was known. These
are the hallmarks that distinguish the mere seeker of geologic
data from the true paleontologist.

The real trick in the field is finding the first fossil in a sedi-
mentary rock. Once this has been collected at the outcrop, the
others come far more readily. Even knowing that years ago
fossils were collected in the general area is a help. If the rock
is a shale that breaks down to a mud and washes away, this first
key fossil may be left as a lag deposit. Crawling on hands and
knees, with nose at ground level, is the time-honored way of
locating it. If the rock is harder, the hammer comes into op-
eration. It may be more poetic to “bring the hammer into
play,” but even when the day is cool and the rock fairly friable,
pounding on an outcrop for extended time periods is hard
work.

The point here is that both of these operations have built-in
limitations on the number of specimens that may be collected
in a short time. The available collecting time at the outcrop,
and the weight one person can easily carry for a short distance,
have been the factors governing the amount of material that
leaves the outcrop. To collect more than three or four bags of
fossils at any one outcrop is unusual. If bulk samples can be
collected rapidly, they are commonly of the type that requires
extensive preparation prior to detailed studies of fossil content.

590. Proceedings of the Biological Society of Washington

Thus, this kind of upper limit also holds for those who study
microfossils. The time involved in taking a channel sample or
digging a trench to collect fresh material may become signifi-
cant. In comparison, for example, with a marine zoologist ac-
customed to collecting on a shallow-water reef, the paleontolo-
gist is a modest collector.

Extrinsic factors important to fossil collecting are not well
understood. It is a general rule that one side of a roadcut will
yield more specimens than another. Whether this is a regienal
feature or whether the phenomenon is related to local factors
such as vegetation, runoff, or microclimate have never been in-
vestigated. Why some fossils in some rocks may be replaced
by other minerals is a major mystery. The conditions that dis-
solve shells but leave their impressions are poorly understood
in detail. The list could be continued.

Intrinsic factors also enter into collecting. There is no sub-
stitute for experience; some rocks just look right for a particular
kind of fossil. In many respects, this is the same as a biologist
knowing the life habitats of a desired living animal or plant
specimen. This type of information can seldom be imparted ex-
cept by word of mouth on the outcrop. At least one attempt
was made to gather these esoteric tidbits as part of general
work on techniques, but the results were far from satisfactory.
The principal point distilled is that collecting is a full-time ac-
tivity. It is possible and often necessary for the paleontologist
to carry on more purely geologic work, such as mapping the
area or measuring the thickness of a rock layer, but these have
to be done before or after the collecting.

There is an interesting minor support of this hypothesis. In
field-work in the western United States, a paleontologist visit-
ing a field man may find more arrowheads in a few days than
the field geologist finds in a season. The geologist strides across
the landscape to get the big picture, but the paleontologist
stays at one spot or shuffles along looking at the ground for his
pet objects. Slow motion is also a fine way to avoid most rattle-
snakes.

One final word should be said about general collecting. It
would be nice if the various disciplines could assist one an-

Natural history collection symposium aieAll

other. In olden days, travel was commonly by train from one
outcrop to another. Because there was time between trains,
some paleontologists used to obtain insects for their colleagues.
It was a nice gesture and one that might be continued even in
these days of more rapid transport, if there was a clear indica-
tion of what other people would like to have collected.

How To Srore Fossi.s

In the how and why of collecting, the why is the easier to
answer, or to at least open the floodgates of rhetoric. Once the
fossils are safely inside a building, the how to store them is far
easier than the why of retention. Compared with other natural-
history objects, fossils are paradise for a curator.

Naturally, catastrophic events may cause serious losses. Type
specimens lost during the great Chicago fire and the flood at
Dayton, Ohio, still cause problems to a few specialists, but,
hopefully, natural-history specimens today are as safe from
such events as might be expected. Good collections are still in
temporary repositories, and undoubtedly a quantity of im-
portant material will be discarded as some universities remove
paleontology from the curriculum, but increasingly the odds
against accidental loss are being lowered.

For convenience, collected natural-history subjects may be
divided into three categories. First, living organisms, which are
stored with great difficulty in zoos and arboreta. Second, re-
cently dead objects, which must be pressed, vermin-protected, or
bottied. Finally, dead things, which do not require watering
and which do not deteriorate. About the only difficulty in pros-
pect for a museum fossil is a coating of the ever pervading
dust. The present-day air-conditioning expert would try to
seduce us into believing that this problem has been solved; it
is better to put one’s faith and one’s specimens in closed cases.

With fossils, one is not troubled by evaporation among
alcoholics, which to the museum-oriented person does not mean
unexplained staff absenteeism. One is not concerned with ma-
terial drying to powder. Except for rare specimens replaced by
pyrite, fossils do not pick up moisture from the air. Fossils are
not edible, and though occasional labels and locality numbers

592 Proceedings of the Biological Society of Washington

may be lost to particularly desperate cockroaches or rats, such
events have been fairly rare in the past and are essentially a
thing of the past. Fossils do not change color after years of
storage, nor do they smell.

About the only obvious and painful drawback to fossil stor-
age is weight. The average collection of fossils, microfossils
excepted, is heavier than the average collection of almost any-
thing else in a museum. One drawer, 28 inches by 22 inches,
full of particularly stony fossils, like colonial corals, requires
complete attention during a moving operation. Drawers of fos-
sils can be stored to a height of 9 feet, but an administrator,
before making a decision for high-level storage, should be
required to carry at least one drawer to the floor. There is a
general rule of nature (Gumperson’s Law) that the heaviest
drawers are always at the top; for any case over 5 feet high this
may become hazardous. It is also well known that museums
that stack drawers rather than place them in cases, keep the
needed specimens in the bottom drawer of a stack (Saunders’
Corollary ).

It is a wise idea to remember always that even though fos-
sils are thoroughly dead, they still retain the ability to move.
When specimens hop from one tray to another, the net result
may be that two otherwise useful collections will have to be
discarded. Trays with deep sides are not a luxury item. Be-
cause it is simply no longer feasible to put locality numbers on
every specimen, stuffing the smaller specimens in glass bottles
has been a technical breakthrough. Clear plastic boxes may
well be worth however much more they cost; if they do come
into general use in the near future, it will be about five decades
since paleontologists stopped putting their prize fossils into
cardboard pillboxes. Folded stand-up labels, in contrast to
those that lie flat, are such a menace to retaining fossils where
they belong and so antediluvian that examples should be put
on special exhibit in the chamber of horrors.

There has been a tendency in unsympathetic administrative
environments to equate storage of dead items with dead stor-
age. If fossils cannot be seen easily, they will not be studied.
Some of the greatest advances that have been made in paleon-

Natural history collection symposium 593

tology stem from some things no more complex than making
aisles wide enough so that drawers may be moved in and out
of cases easily. Lighting adequate to permit specimen examina-
tion in a storage area has done more for overall clarification of
species problems than the most sophisticated hardware of biom-
etry.

Wuy Do WE Botuer To Keep Fossis?

There are so many reasons not to keep collections that one
hesitates to open this question for discussion. Collections take
up space, and space is money. They take up time, and time is
money. About the only reason for keeping them is for the sake
of honesty. If less painful words are needed, collections are
kept for purposes of documentation and scientific verification,
as well as to provide raw material for new studies. The Wash-
ington, D. C., collection includes more specimens that should
be retained for purposes of biologic and geologic documenta-
tion than any other in America. There may be some merit in
the view that once an optimal or critical size is reached, the
importance of a collection increases more rapidly than its bulk.

A gifted mathematician may derive four from two plus two.
Once this is published, another specialist with the proper com-
putation can verify this discovery. In marked contrast, a rela-
tive date based on a fossil occurrence or a biological descrip-
tion of it is not nearly so tidy. No matter how good the printed
description or how accurate the figures, sooner or later they
are found wanting. If a paleontologist is smart, he will never
completely trust the published work of another, but will look at
the specimens in question. If he is particularly intelligent, he
will not even trust his own published work and will continue to
reexamine his fossils.

Systematic biology is an additive science and does not make
great strides forward to major unifying natural laws. It does
not lend itself to the sporadic quantumlike great leaps forward
that have characterized the history of the physical sciences.
Like all other kinds of systematics, paleontology moves for-
ward at a crawl, building its monumental truths a dust particle
ata time. We will probably never know with the precision of
a mathematician the absolute stratigraphic range or total bio-

594 Proceedings of the Biological Society of Washington

logic diversity of a single extinct species, let alone the millions
of such species that are in various stages of study, from those
still awaiting collection on the outcrop to those in the latest
published monograph.

However, every bit of new information throws a faint glim-
mer onto the overall biologic-stratigraphic system, and old
material ought to be reexamined in this light, no matter how
feeble the light may be. The great weight of fossil specimens
described and those yet to be described is good ballast to keep
the hot-air balloon of theory from rising too high. For the
paleontologist, particularly, one battle cry is alpha taxonomy
forever! For this sort of old-fashioned work, one needs to look
at specimens.

If one agrees that material should be kept, the logical posi-
tion is to store it in the most useful system. This presumes a
purpose in study, but the true paleontologist really has two
purposes. One is biology and the other is stratigraphy. As a
consequence, varying shades of schizophrenia infect the col-
lections. In Washington, Geological Survey collections are
stored in stratigraphic order and National Museum collections
are stored partly in stratigraphic order, but mainly in biologic
order. Types are stored in alphabetical order, for “conveni-
ence,” an infelicitous expression if there ever was one. This
dual system is found at most institutions that retain fossils. Of
course, the outsider immediately objects that a unique speci-
men cannot be in two systems at the same time. This is ab-
solutely true, but the dual system still works somehow and is
used in most major collections throughout the world. Once the
details of a particular local arrangement are understood in a
museum, the paleontologist readily pursues his specimens up,
down, and sideways through the collection.

The precise arrangement of the individual lots within a
stratigraphic or biologic series is a subject for violent argument.
One quick way to provoke argument is to state complete op-
position to any arrangement by numerical sequence, for this is
a simple method to follow; such simplicity is a trap. Collections
should be in a subject matter arrangement just exactly the same
way books are arranged in a library. Often the particular bit

' Editor’s italics—author’s exclamation point.

Natural history collection symposium 595

of information desired lies in the adjacent collection, just as
the book you finally choose is adjacent to the one you origi-
nally thought you wanted.

A few words should be said about mechanics, because a
poorly kept collection is a powerful administrative argument
for discarding all fossils. In the part of the Washington mega-
fossil collection that seems in best arrangement, the crucial ele-
ment in the system is the one person whose job it is to keep
track of things. About 40,000 collections are involved in this
70 year accumulation. Given a locality number, a particular
lot may be located in 2 to 3 minutes. Because there is a logical
arrangement, a blind search for fossil data on a restricted age
and area basis can be run in less than half an hour. The only
trouble with the system is inadequate manpower to bring all
collections into proper curatorial shape within the system. We
can keep current, more or less, but the backlog from past years
is not reduced.

Automatic data processing will not help one iota in typing
locality descriptions or preparing specimens. Some persons as-
sume that an old system is necessarily outdated, whereas a
more correct assumption is that the system has been time-
tested and found to be successful. The classical methods have
been “debugged,” to use the current argot. Changing them may
not be a wise investment of time or money.

The storage situation may be a bit more complicated with
microfossils because one cannot simply look at the specimens
with a hand lens. However, the same general principle holds,
in that the collections should be arranged in a logical order.
The nomenclatural situation within the field of foraminiferal
studies is chaotic and is expected to get worse. The one reed
left to cling to is the system of filing microfossil slides in al-
phabetical order by the original name. It works. Other kinds
of microfossils may be filed by other arrangements.

This leads to the conclusion that the best system for any
institution to follow is that which satisfies the workers most
closely concerned. If this sounds trite, silly, and obvious, the
other side of the coin is that an institution should be willing to
stand the expense of major reshuffling as workers and ideas

596 Proceedings of the Biological Society of Washington

change. Libraries reshelve books when necessary and survive
the process. Paleontologists generally are too xenophobic and
ergophobic to put collections in the order that yields maximum
information for their own purposes.

Granted that all published or cited material should be kept,
something should be said about the residue. Many institutions,
but particularly universities, tend to hang onto material too
long. Much rock gathered during the preparation of a thesis
should be discarded; the good material should be properly
curated and saved. Junk brought in decades ago by field men
and never cited can be discarded. Fossils do not age materially,
but accompanying data may become obsolete. A collection “Car-
boniferous, Indian Territory” was important last century, but
its time of significance is long past. To give another example,
the push today is in paleoecology, but the collections made by
prior generations are too biased to yield automatically the new
data needed without additional field investigations. Field in-
vestigations in any area of natural history, including paleontol-
ogy, always seem to yield collections!

Although it is easy to say discard unnecessary material, it is
most difficult to do. One general rule to follow is that no one
under 40 should be permitted to discard collections gathered
by earlier workers. Often biologically poor material may be
stratigraphically important and vice versa. Unless one has done
fieldwork in rocks of a particular age and area, the best course
is keep all the material already available for that age and area
in storage. It is far better to err on the side of keeping too much
than to discard an unmarked type specimen.

The time to discard is before collections are given numbers.
Inadequate collections should be promptly abandoned and
not left in odd corners, following the current method of con-
tinuing the sins of our predecessors. Proper curation is a
thankless task which is generally shirked. Shame on all of us.
If a fossil is worth keeping, it is worth keeping well.

Much as one hates to weaken a particular point, I must ad-
mit that although there are many good reasons for consolidat-
ing and discarding collections, economy is not one of them.
One of the most expensive operations is to selectively prune

Natural history collection symposium 597

collections. It is possible to work for a year and empty one or
two storage cases.

Unless there is someone who has adequate time and cares
enough to put the fossils in some order, all that results from
collecting is a random arrangement of limited value. At the
risk of annoying people further, a minor semantic needle
should be emplaced. In my ancient Funk and Wagnalls, the
word “collection” implies unorganized and promiscuous char-
acter similar to that of assemblage. This is hairsplitting, but it
just could be that parts of our collection are properly so desig-
nated.

THE FUTURE

Scientists are supposed to make predictions, probably to
prove that they are human and can be as mistaken as anyone
else. Long-range predictions are better to make because the
audience to whom the prediction was made is no longer around
to ask questions. The alternative and next best method, which
is followed here, is to make conflicting predictions, so that one
prediction of the two may prove right.

Growth rates of collection bulk might be meaningful. By
averaging a sample of palynologists, coral specialists, elephant
hunters, and other assorted paleontologists, I have arrived at a
figure of three museum cases 3 feet high per year. As these
cases occupy 6.6 square feet and are usually stacked two high,
space may be used ata rate of 10 square feet per year, plus all-
important aisle space. Fifteen square feet of growth per man
year is an authoritative wild guess. Thus, the new paleontolo-
gist starting out should be assured of 450 feet of space to fill
with his collections, not counting what he will inherit in his
specialty.

Unfortunately, collections simply do not grow this way. A
better comparison is with growth studies of fish. If a large
number of infant minnows are crowded into a small tank, they
are stunted. When these stunted fish are transferred to a
larger aquarium, however, they immediately grow to normal
size. Available space determines the size of collections, not
vice versa. Paleontologists assigned to new quarters with fresh
storage space fill it rapidly and then are cramped until the next

598 Proceedings of the Biological Society of Washington

building provides a quantum jump. This principle has been
checked at several localities and holds for at least North
America and Europe.

It is also safe to predict that no extensive buildings for pa-
leontologists in these regions are anywhere obvious on the
horizon, Even more important, administrators have not been
trained to think of large collections as scientific instruments.
Major advances in other fields are accompanied by major in-
vestments in hardware. Probably the same principle applies in
paleontology. Cyclotrons, sounding rockets, and radio tele-
scopes really are not that different from new buildings filled
with old organisms. Larger collections and advances in the
field go hand in hand.

Another consideration beside storage space to fill is source
of fossils. Most fossils gathered to date have been the product
of long-time weathering processes. Once specimens are picked
up from the outcrop surface, years of weathering are needed
before others may be released. Some conservation-minded pro-
fessors have preserved favorite outcrops only by extorting from
a class all fossils collected and then sprinkling them back on
the outcrop for next year’s crop of budding experts to find.
Most classic localities in this country have been picked or
hammered clean of specimens.

Worse still, new exposures are not being developed for fos-
sils. Lots of fossils once came from limestone quarries, obtained
by the workers who were crushing stone by hand. Today, the
rock is untouched by human hands from quarry face to cement
bag. Railroad cuts used to be wonderful places to find fossils,
but is there anyone still alive who can remember the last time
a new railroad line was laid out. Highway cuts ought to be
fine for collecting and were so for many years. No one is op-
posed to major erosion control, but the highway engineers think
of erosion the same way as prohibitionists think of alcohol and
consider even a tiny amount sinful. To see grass being sown on
potentially highly fossiliferous roadcuts before even the con-
crete slab is poured is most discouraging. It is fairly safe to
state that the bulk of the fossils that can be obtained easily from
the weathered crust of the United States have been obtained
and are stored away.

Natural history collection symposium 599

Having demonstrated why collections will not markedly in-
crease in size, let me now take the counter argument. The wave
of the future is already upon us, without any plans for coping
with it. Paleontologists have known for hundreds of years that
some fossils have been replaced by minerals that are insoluble
in certain acids. Because of this, some outcrops have yielded
choice fossils, or a specimen might be cleaned with a tooth-
brush soaked in acid, or one or two specimens might be freed
from the rock matrix by placing the matrix in an acid-filled
beaker.

Three decades ago, one of the senior National Museum
paleontologists noted that chemical change of fossils persisted
through the thickness of the rock. This fact was not new; more
than half a century ago, fossil corals were dissolved from rocks
and sold. However, he put an entire limestone block in acid,
and then another, and another, and another. ... The results
have shaken the paleontologic world. The specimens obtained
have been strikingly beautiful and highly significant both bio-
logically and stratigraphically.

Perhaps even more significant for this discussion is the bulk
of silicified fossils. By spending the same time at the outcrop,
collecting limestone blocks rather than loose fossils, the number
of specimens increases by many orders of magnitude. One
hundred good specimens of a species from a single locality has
been exceptional in the past. Now, a number of species are
known from an entire case of choice material.

Silicified fossils are not sturdy. We have leaped from stor-
ing rocks to storing objects as delicate as butterflies. One does
not pile up a heap of silicified fossils in the corner of a drawer.
Good ones should be chemically hardened. They ought to be
stored on cotton and even packaged individually. They have
to be protected from the sudden jerk and slam of the conven-
tional drawer with the sticking runners. When these fossils
were first shipped between museums 20 years ago, the only
known method was to imbed them in wax, and some have never
been cleaned free of it. It has taken years just to stumble on the
obvious idea of shipping them packed in sawdust. The field
is wide open for new techniques.

600 Proceedings of the Biological Society of Washington

All these new factors, brought into the picture by silicified
fossils, mean a tremendous increase in space; I have no esti-
mates other than “lots more.” If field funds, preparation facili-
ties, and technical assistance were optimum and permitted
paleontologists to really move into the silicified fossil business
in a businesslike way, the entire character of the collections
could be changed in two decades.

Methods employed in obtaining silicified megafossils do
not work for all paleontologists. Certainly, those who work on
microsfossils and micro-microfossils should not be slighted, but
one seldom has thought of them as requiring a great deal of
space. However, new chemical and mechanical techniques
have demonstrated that fossils are to be found in almost all
sedimentary rocks. Today, it is a question whether the micro-
scope slides or the black boxes and cameras take up more area.
Suddenly the micropaleontologist wants a great deal more
from life than space for a one burner stove to boil his pot of
mud and a desk drawer to store slides. He may never indivi-
dually require as much space as the bulldozer-wielding whale
collector, but there are many more of the people working on
the little bugs. Curiously enough, when prepared residues are
retained along with their fossil content, more space is needed;
no one throws away residues because there may be a need
later to search in them for more microfossils.

This leads me to the final set of summary predictions. We
will need substantially new buildings and much better hand-
ling and storing techniques for silicified fossils. Probably the
best method will be to organize two separate collections, based
entirely on the mechanical strength of the fossils. As a parallel
development, microfossils are amenable to an organized, fully
automated specimen storage and retrieval system. The pa-
leontologist need only punch a few buttons on his room con-
sole to have the necessary slides moved into his microscope
field.

For a century and a half, fossil storage has been essentially
unchanged. Twenty years from now it will be all different. I
have no idea where the money for a major national investment
in paleontology will be obtained. The physicists became fat off

Natural history collection symposium 601

of radar, and the chemists have done fairly well as a result of
the atomic bomb. If, as a result of the moon race, the first ex-
traterrestrial hand sample is fossiliterous,' perhaps paleontology
will also reach the land of cornucopia. Until that golden day,
collect new old fossils and keep them, no matter how tight the
quarters, for only in this way will life continue to flow in the
dry bones.

! Unfortunately, it wasn’t and further the photographs of Mars do not look par-
ticularly promising as a nice place to visit, let alone live. However, there is always
the hope of Venus or litter dropped by UFO’s.

602. Proceedings of the Biological Society of Washington

THE ROLE OF THE NATIONAL PARASITE COLLEC-
TION IN VETERINARY PARASITOLOGY

By Wittarp W. BECKLUND
Beltsville Parasitological Laboratory
Animal Disease and Parasite Research Division
Agricultural Research Service
U.S. Department of Agriculture, Beltsville, Maryland

Almost every individual during his lifetime has at least once
become a collector of objects purely from curiosity or for the
love to determine the quantity and variety of objects which
can be accumulated. Naturally, pride is taken in the collection
and in its exhibition. This is probably the way natural history
collections first began, and no doubt many individuals still
think of them in this light. Many collections, however, now
constitute working tools in research as essential as are the ex-
perimental animals or laboratory equipment used. Among
these is the parasite collection herein discussed. It was primar-
ily started, and is still used today, to determine what parasites
cause disease, their geographic distribution and animal hosts,
and the diagnostic characters by which the parasites may be
identified, including their various immature forms. This in-
formation is essential for treatment, control, quarantine, and
research purposes. The collection is described herein, along
with an index to the literature on parasites with which it is
used, and some examples are given of its current and future
roles as a working tool in veterinary parasitology.

PARASITE COLLECTION AND INDEX To PARASITOLOGICAL
LITERATURE

Animal disease workers in the U. S. Department of Agricul-
ture recognized the need for a parasite collection and index to

50—Proc. Brot. Soc. WasH., Vou. 82, 1969 (603 )

604 Proceedings of the Biological Society of Washington

the literature on parasites over 76 years ago when they estab-
lished the National Parasite Collection and the Index-Cata-
logue of Medical and Veterinary Zoology. The latter is a com-
pendium of the world’s literature on parasitology. Both of these
working tools are maintained at the Beltsville Parasitological
Laboratory where they are used by the some forty scientists at
the Laboratory, as well as many visiting scientists. The records
of the Collection enable investigators to find quickly essential
information on the parasites deposited therein; the Index-
Catalogue serves the same purpose with respect to the world’s
literature on parasites. The history of the Collection and a
detailed description of the various parts and publications of
the Catalogue are given elsewhere (Becklund, 1969a, b).

The Index-Catalogue consists of an Author Catalogue, and
four Parasite-Subject Catalogues, namely: Parasites (sub-
divided by taxonomic groups), Hosts, Subject Headings (e.g.,
biochemistry, cultures, immunology, etc.), and Treatment. All
information is recorded on 3” x 5” cards which are filed in ap-
proximately 1500 drawers. Over 100 publications comprising
more than 20,000 pages have been issued under the title Index-
Catalogue of Medical and Veterinary Zoology.

The Collection is composed of parasitic protozoans, cestodes,
trematodes, nematodes, pentastomes, lice, mites, ticks, and
other miscellaneous parasites. Approximately 65,000 lots, con-
sisting of one to many specimens each, have been accessioned.
Most of the specimens were collected during research and
regulatory activities of the U. S. Department of Agriculture
and state animal disease agencies. The procedure for the de-
posit of specimens in the Collection, so that they, and all in-
formation about them, can be easily found, is as follows:

1. All pertinent information on the name of the parasite, host,
location in or on the host, geographic locality in which col-
lected, and collector's and identifier’s names and dates are re-
corded on accession numbered (Collection number ) forms; this
information is also recorded by Collection number in books
and on labels that are placed with the specimens. 2. Each lot of
specimens is assigned a storage number which designates its lo-
‘ation in the Collection, and this number is recorded with the

Natural history collection symposium 605

aforementioned information. (Most liquid preserved material
is stored in two-ounce square bottles in numbered wooden
racks holding several bottles. Slides are stored in numbered
wooden boxes holding 25 slides each.) 3. A Parasite Index and
a Host Index are maintained on cards by recording, for each
lot of specimens, the name of the host, Collection number, and
storage location number under the name of the parasite in the
Parasite Index; and the name of the parasite, Collection num-
ber, and storage location number under the name of the host
in the Host Index.

CurrRENT Uses Or THE PARASITE COLLECTION IN
VETERINARY PARASITOLOGY

The Parasite Collection is a depository for specimens that
have been mentioned in published reports. Specimens desig-
nated by authors as type material, as well as those that represent
new host and distribution records or various forms in the life
cycle of a species, are regularly deposited. In addition, the
Collection and its records serve scientists in various ways. The
following are a few examples:

Description of new species from man and animals: The thou-
sands of authoritatively identified specimens in the Collection
are invaluable for comparative purposes to determine species
that are new to science. Several hundred new species of para-
sites have been described by U.S.D.A. scientists. During the
last five years the Laboratory's personnel have described new
species from various hosts, including man, marmoset, mountain
goat, deer, alpaca, vicuna, and an African lizard. Among these,
the species described from man is particularly important. It
has reportedly caused over 900 cases of human capillariasis in
the Philippines, some of which were fatal (Chitwood, et al.,
1968).

Provide information on the parasites of domestic animals in
North America: A checklist of parasites of domestic animals in
the United States and Possessions, and Canada, was prepared
in 1945 (Dikmans) and revised in 1964 (Becklund). It was
prepared for use in teaching, in research, and in regulatory
quarantine activities from information in the Collection records

606 Proceedings of the Biological Society of Washington

and Index-Catalogue, and by examining many specimens on
deposit. This checklist gives the common and scientific names
of the parasites, location in or on the host, intermediate hosts in
the life cycle, if any, and geographical distribution. One hun-
dred and twenty-one species of parasites were listed from cat-
tle, 119 from sheep and goats, 92 from equines, 72 from swine,
132 from dogs and cats, and 179 from chickens, turkeys, pi-
geons, pheasants, ducks, and geese.

Provide information on ticks of veterinary importance on
imported animals and items: For many years ticks that
were removed from domestic animals, exotic wild animals,
and items offered for entry and imported into the United States
were identified and deposited in the Collection. Many of the
ticks are of medical and veterinary importance; therefore, the
species, hosts or items on which they were found, origin, and
locality where collected, were recorded for regulatory and re-
search purposes (Becklund, 1968). The ticks were removed
from cattle, horses, numerous kinds of zoo animals ranging
from hedgehogs to elephants, beef, cattle hides, palm leaves,
mailbags, medicinal herbs, bird guano, and hair, from many
parts of the world. This study revealed that: (1) The ticks rep-
resented nine genera and 37 species; (2) most of the exotic
ones were males; females apparently drop off at foreign quaran-
tine stations and while the imports are en route; (3) native as
well as exotic ticks on imported animals can be vectors of exotic
diseases; and (4) harmful ticks can occur on unexpected strange
items and abnormal hosts.

Provide specimens to establish differential characters to dis-
tinguish between species: American sheep are hosts to several
species of thread-necked strongyles. The various species are
very similar, morphologically, and have been confused with
one another for more than 70 years. Reports of the pathogenic-
ity, treatment, incidence, hosts, etc., of the various ones are
therefore questionable. A study of hundreds of specimens of
these worms comprising 90 lots of specimens in the Collection
revealed that instead of two common species and three rare
ones, American sheep are parasitized by three common species
and three rare ones. This study of specimens enabled investiga-

Natural history collection symposium 607

tors (Becklund and Walker, 1967) to establish morphologic
characters to readily identify the species and determine their
geographic distribution. Subsequent work with additional spec-
imens from the Collection (Stringfellow, 1968) revealed a
hitherto unrecognized structure of the worms that is useful in
their identification.

Provide information on the probable transmission of para-
sites between domestic and wild animals: Parasites in the Col-
lection and information in the Index-Catalogue were used to
supplement findings from a study of the parasites found in
18 bighorn mountain sheep in Montana (Becklund and Sen-
ger, 1967). The known number of bighom sheep parasites was
increased from 34 to 51 species. Thirty-six of these 51 species
are known parasites of domestic sheep and 18 parasitize cattle
in North America. Thus, in regions where these animals graze
on the same range land, parasites are probably interchanged
between bighorn sheep and domestic sheep and cattle.

Provide specimens used to evaluate malformations in para-
sites resulting from an antiparasitic chemical: At necropsy,
sheep suffering from haemonchosis and receiving therapeutic
doses of phenothiazine had numerous deformed male speci-
mens of the large stomach worm (Becklund, 1960). The per-
centage of deformed male worms in populations exposed to the
drug ranged from 0 to 47 percent, whereas the percentage in
unexposed populations, many of which were obtained from
those placed in the Collection from 1900 through 1939, before
the advent of phenothiazine, ranged from 0 to 0.3 percent. The
parts of the worms affected, because of their prominence and
normally characteristic conformation, are used in systematics;
consequently the antiparasitic somewhat weakens current com-
petence to distinguish species. In some respects, the deformi-
ties studied are similar to those recently reported in human
medicine involving thalidomide.

Future Uses Or THE PARASITE COLLECTION IN
VETERINARY PARASITOLOGY

Parasitologists hope to rear parasites through their entire
life cycle without the host (in vitro cultivation). Great strides

608 Proceedings of the Biological Society of Washington

have been made recently along this line of research and nema-
todes of domestic ruminants have been reared in vitro from the
eggs to adults (Leland, 1967). Parasites so reared are unique
and should be compared with specimens from animals to estab-
lish their normality before their cultivation is considered a
complete success. Because of their uniqueness, and the pos-
sibility of correlating their structure and development with
nutritional deficiencies in the culture media, representative
specimens have a place in the future of the Collection. This
was recognized by Schiller (1965), who has deposited in vitro
reared specimens and indicated their Collection number in his
report.

Events of the past strongly suggest that the Collection will
continue to be an essential working tool. Specimens will prob-
ably be needed for the evaluation of any change in morphol-
ogy, pathogenicity, or host of parasites. Such changes could
result from research activities, such as the use of X-irradiated
larvae to produce immunity in animals, or from adverse en-
vironmental conditions which affect the host or the parasite,
such as pollution, pesticides, or radiation. Hence, a problem
today is deciding what kind and how many specimens are
needed to fulfill future needs.

LITERATURE CITED

BreckLuNb, W. W. 1960. Morphological anomalies in male Haemonchus
contortus (Rudolphi, 1803) Cobb, 1898 (Nematoda: Trich-
ostrongylidae ) from sheep. Proc. Helm. Soc. Wash., 27(2):
194-199.

1964. Revised check list of internal and external parasites of
domestic animals in the United States and Possessions and
Canada. Am. J. Vet. Res., 25( 108): 1380-1416.

1968. Ticks of veterinary significance found on imports in
the United States. J. Parasit., 54(3): 622-628.

1969a. National Parasite Collection at the Beltsville Parasito-
logical Laboratory. J. Parasit., 55(2): 375-380.

1969b. Index-Catalogue of Medical and Veterinary Zoology
at the Beltsville Parasitological Laboratory. J. Parasit., 55(2):
381-384.

AND C. M. SENGER. 1967. Parasites of Ovis canadensis cana-
densis in Montana, with a checklist of the internal and ex-
ternal parasites of the Rocky Mountain bighorn sheep in
North America. J. Parasit., 53(1): 157-165.

Natural history collection symposium 609

AND M. L. WALKER. 1967. Nematodirus of domestic sheep,
Ovis aries, in the United States with a key to the species. J.
Parasit., 53(4): 777-781.

Cuitwoop, M. B., C. VELASQUEZ, AND N, G. SALAzAR. 1968. Capillaria
philippinensis sp. n. (Nematoda: Trichinellida) from the
intestine of man in the Philippines. J. Parasit., 54(2): 368-
Srl

Dixmans, G. 1945. Check list of the internal and external animal para-
sites of domestic animals in North America. Am. J. Vet. Res.,
6(21): 211-241.

LELAND, S. E. 1967. In vitro cultivation of Cooperia punctata from egg
to egg. J. Parasit., 53(5): 1057-1060.

ScHILLer, E. L. 1965. A simplified method for the in vitro cultivation of
the rat tapeworm, Hymenolepis diminuta. J. Parasit., 51(4):
516-518.

STRINGFELLOW, F. 1968. Bursal bosses as a diagnostic character in Nem-
atodirus of domestic sheep, Ovis aries, in the United States.
J. Parasit., 54(5): 891-895.

610 Proceedings of the Biological Society of Washington

THE NATIONAL COLLECTIONS AS BIOLOGICAL
STANDARDS

By RicHarp COwAN
Smithsonian Institution, Washington, D. C.

Gathering of natural history objects must be as old as man
himself and a reflection of his inherent curiosity in the world
about him. These objects, at first, must have had primarily
utilitarian interest, having some real or suspected property of
direct survival advantage. Eventually we can imagine that cer-
tain aesthetic properties such as color and form began to be
important. Later, as he became ever more sophisticated and
less concerned with daily survival, such objects were gathered
in “cabinets of Curiosities’——pretty stones, fossil bones,
brightly colored butterflies, etc. They may have served much
the same purpose as modern coffee table picturebooks—as
conversation pieces. Ultimately, specimens of the natural world
were recognized as important documentation of the kinds of
organisms, their geographic distribution, their variability, and
their evolutionary history. Systematic collections as biological
standards began with that realization, and, with the literature
their study has generated, they are still the basic tools of the
systematic biologist.

The natural history collections of the U.S. National Museum
had a very early origin in the enormous collections brought to
the Smithsonian by Spencer Fullerton Baird and added to by
virtually every serious biologist since. Increasing by about one
million specimens annually, they now total somewhere be-
tween 50 and 60 million. One cannot speak of the growth of
this major scientific resource without acknowledging the very
large contributions to the National Collections made by the
Geological Survey, the Fish and Wildlife Service, and the

51—Proc. Bion. Soc. WasuH., Vor. 82, 1969 (611)

612 Proceedings of the Biological Society of Washington

Department of Agriculture entomologists, by whatever titles
these groups may have been known earlier. While the care of
the collections is the legislated responsibility of the Smithso-
nian Institution, they are what they are because of many, many
years of cooperative development.

While these National Collections have grown both qualita-
tively and quantitatively and provide an almost unparalleled re-
search resource, it can be said that we have not yet reached
maturity in one important aspect. Although we often receive
type materials and important sets of material documenting a
particular study, we have not achieved the stature in this
country that the British Museum has achieved in Britain, where
to have one’s collections incorporated is a mark of scientific
distinction. Rather than relying on legislation, we must dem-
onstrate our willingness, even eagerness, to serve as the Na-
tion’s repository of biological standards, which, like physical
standards, must be preserved at a site that has a reasonable
chance of caring for them in perpetuity.

It might be well at this point to consider the question of who
uses these standards and for what purposes, especially in view
of the increasing costs in time, space, and dollars to maintain
them. The collections are used constantly by systematists in uni-
versities (many of whom have disposed of such collections), as
well as those in other museums. Last year (1967) we sent 372,886
lots and/or specimens to other researchers over the world. In
addition, we hosted 1,195 student or professional research biol-
ogists who spent 7,003 man-days in our museum. Most of this
sort of use is obviously a service to the systematic community
but others use the standards as well. After the Pacific testing of
nuclear devices, concern developed in many quarters about
radioactive contamination of the environment, especially of res-
ident plants and animals. But how could anyone guess what
the condition of the biota was before the tests? Specimens in
the National Collections from early expeditions in the test area
provided the answer to that question—a biological standard
provided the basis for solving this important problem. Other
examples of the use of these standards are plentiful. I wonder
if the historian considering the development of American cul-

Natural history collection symposium 613

ture can be really thorough without an understanding of the
role of the undisturbed biota on which the colonists depended
and with which they contended. How can we talk about re-
storing the quality of the environment without referring to
these standards to learn what lived where and when? Another
less obvious application of biological standards, that is collec-
tions, is in understanding such dramatic, evolutionary explo-
sions as occurred in Rubus, the blackberry genus. Before the
development of agriculture in the eastern half of the country,
the species of this genus were nicely separated from each other
by ecological and geographic factors of one sort or the other,
but as the forests were leveled to make farm land, new oppor-
tunities opened up for once-separate species to commingle ge-
netically and the result has been chaotic for the systematic bot-
anist. His understanding of the environmental situation in the
earliest part of the history of this region illuminates the sub-
sequent man-made confusion. The list of examples could be
very long, but I doubt that anyone here, at least, will question
the importance and value of the National Collections, or that
they are used. At this point, f should like to mention the ob-
vious, that the collections to be valuable for future problem-
solving must be housed, cared for, and added to—and these
present real, very difficult problems.

One of the most critical has always been that of space for
housing collections. Growth of collections, even under normal
circumstances, is difficult because of space and financial limita-
tions, but we are at this moment entering a period of unparal-
leled expansion of various types of field biology. When the
International Biological Program and the numerous large, fed-
erally-supported environmental studies get underway, the enor-
mity of the problem of caring for the mountains of documen-
tary collections that surely should result staggers the imagination.
All of us, to varying degrees, will be faced with the problem
of how to process these materials so that they are available to
biologists generally, systematists, physiologists, ecologists, and
perhaps even the molecular types as well. As the numbers of
collections grow, there is increasing difficulty with even bring-
ing together the existing specimens of a particular group, and

614 Proceedings of the Biological Society of Washington

an even more formidable task of gathering and synthesizing
the data attached to the specimens. The mundane problem of
housing and caring for these constantly expanding collections
poses serious space and time-use problems requiring our most
serious consideration of the quality, the nature, and the meth-
ods of curating the collections. At one point, J questioned that
very much thought was given to what is added to the National
Collections, for | am sure we can all agree that undisciplined
growth is detrimental to their long-term usefulness. Within
the past year we have begun to write what may be termed a
rationale for collections growth, and I have been pleased to
see numerous examples of correspondence that indicate real
judgment on the part of the curators in rejecting substantial
collections. In earlier times of our history, as well as that of
other collections centers, there may have been more justifica-
tion for considering the largest collection the most important
but the attention given to qualitative considerations is very
important at this point in our history.

Aside from being more selective in adding to the National
Collections than at times in the past, how can we solve, or at
least ameliorate, the problem of space for collections? One ap-
proach is to give serious thought to the nature of the materials
we maintain. Why should each systematics center strive for
world-wide, in-depth coverage of all groups of organisms? Isn’t
it possible to think of an organized sharing of the responsibility
of developing the degree of coverage required by the needs of
biological research? There is precedent for this. Twenty or 30
years ago, several of the large systematic botany centers, all
with deep interests in Latin American plants, got together and
agreed to divide the job of developing tropical plant collec-
tions. Each center concentrated on collecting and studying the
plants of a single country or region. In addition, each institu-
tion shared representative collections from their special regions
with all the others of this informal consortium. The plan
worked remarkably well and to some extent it is still observed
by the participants. Perhaps the cooperation achieved in that
instance could serve as a model for broad consideration of col-
lections-space problems.

Natural history collection symposium 615

Another way of looking at the problem, one that has been
suggested previously, is that of inter-institutional transfer of
blocks of collections on a long-term loan basis when the borrow-
ing institution has a specialist not represented on the staff of
the loaning institution. It is perhaps unnecessary to state the
obvious, that there is no center in existence that can hope to
employ a specialist for each of even the largest groups of orga-
nisms. Could a collection not under active study by a specialist
at one institution be housed with a specialist at another? To do
this, we would have to develop common curatorial standards
that would ensure that the collections of the one institution
were cared for equally well by the borrower. We often assume
this for present-day, smaller loans and sometimes are dis-
appointed but surely we could determine the standards for
specimen cases, the kind and frequency of application of fumi-
gants, and the sort of fire-protection required for preserving
each other's collections.

A second major problem of the National Collections, a prob-
lem shared with all other Federal systematic centers, is that of
grossly inadequate supportive assistance—technicians, aids,
research assistants and the like. For the past ten months I have
chaired an interagency panel charged with a consideration of
the state of health of systematics in the Federal system—some
of the panel members are surely in this meeting. We learned
that the average level of support is about one supportive person
to each professional which is about 30 percent of what has been
recommended as adequate for scientists in Federal laborato-
ries. It can scarcely be denied that employing well-trained, ex-
perienced scientists and then using substantial parts of their
time in non-scientific tasks is the most absurd sort of ineffi-
ciency. These problems of space for the collections and the cura-
torial assistance to manage them must be solved if the National
Collections are to continue to be useful biological standards in
the future.

One of the most important developments for systematic biol-
ogy is that of data processing technology as it can be brought
to bear on repetitive, non-scientific chores. Efforts are being
made, mostly at the pilot-project level, by several museums to

616 Proceedings of the Biological Society of Washington

assemble the data associated with some collections in a ma-
chine-retrievable form. If one assumes even ten facts in associa-
tion with each of our 50 million specimens, it is obvious why
progress in systematic biology is slow but it also suggests that
the task of computerizing even major parts of such an enor-
mous data-base requires very careful planning and decision-mak-
ing. Machines can handle the problem of cataloging and re-
trieving published data as well, but the annual exponential
growth both in collections and literature makes action increas-
ingly urgent. For data-processing applications to have the great-
est usefulness, cooperative data-banks based on inter-institu-
tional agreement will be important. To achieve this cooperation
there should be some agreement about what information will
be deposited in the bank to answer what sort of questions. The
expense of the automatic data-processing operation is such that
the bank should neither contain trivial information nor be
queried for it. While it is imperative that we develop a com-
mon system, or at least compatible ones, the provincialism of
many of us seems to indicate that this will be one of the major
problems that may be solved for us by the funding sources and
the computer hardware people. In this respect, we need a
common approach among the principal natural history mu-
seums such as the New York art museum consortium has
evolved; a united viewpoint still breeds confidence and _ at-
tracts the support of others. At the same time we are attempt-
ing to develop national cooperation, we need to consider how
we can work closely with major collections centers in other
parts of the world. Free access of systematic information is
necessary for the maintenance of the position of systematic
biology and closely allied biological disciplines as primary con-
tributors to science.

As we have heard from some of the preceding speakers, a
beginning has been made in the area of recording information
associated with new collections and to some extent with the
older collections as well. While it may well be impractical to
think of computerizing the data on all 50 million collections,
this surely should not discourage us from storing data at some
appropriate level and in some instances to the specimen level.

Natural history collection symposium 617

I think we must face the fact that one of the most substantial
problems in the area of data-handling is ourselves. Our gen-
erally narrow specialties often lead us into a sort of scientific
isolationism, an inwardly directed concern for our own in-
terests. We are often constrained by a traditional mode of op-
erating, which we feel uncomfortable about discarding or mod-
ifying. Consequently, as we face the increasingly critical need
to recover data from collections and associated literature, we
may respond by burrowing more deeply in our traditional
methods of data-gathering and data-handling with consequent
loss of time for a function that is not always recognized as part
of the systematic job—interpretation of the data we gather and
organize. The Museum of Natural History, with the strong
backing of the administration above, is seeking appropriated
funds for carrying out the kinds of data-processing applications
that will make the information in the National Collections more
available to the entire scientific community. The pilot pro-
grams now current in the museum, supported by the HEW
contract, is an effort in which we can all share the leadership
role that is so appropriate for those of us associated with these
Collections. It is not an effort of one person or even of a small
group of curators, but rather a means of getting started toward
the long-range goal of making the collections more significant
for ourselves and for our colleagues, many of whom expect us
to provide such leadership.

If the National Collections are biological standards, then we
who are the keepers must be prepared to lead, to discard the
traditional when it no longer meets needs, for if the standards
fail to provide the information needed to solve problems, they
will cease to have importance to anyone but ourselves.

618 Proceedings of the Biological Society of Washington

DOES ANTHROPOLOGY NEED MUSEUMS?

By WILLIAM C. STURTEVANT
Smithsonian Institution, Washington, D. C.

You can be a museum, or you can be modern, but you can’t be both.
Gertrude Stein (refusing to leave her collection to the Museum of Mod-
ern Art).

Rien ne me parait ressembler autant 4 un bordel qu’un musée. On y
trouve le méme cété louche et le méme cété pétrifié. . . . Dans Pun et
Vautre endroit on est, d'une certaine maniére, sous le signe de l’archéolo-

gie; et si j'ai aimé longtemps le bordel c’est parce quwil participe lui aussi
de l'antiquité, en raison de son coté marché desclaves, prostitution rit-
uelle.—Michel Leiris, Chargé de Département d’Afrique Noire, Musée
de THomme (1939: 41).

Museum anthropologists often bewail the present state of
anthropology in museums, not infrequently blaming this on a
wrong turning taken by some of the most prestigious areas of
anthropology a few decades ago. If only the leaders of our
field could be brought to recognize their mistakes, they would
again send their students to museums and the Golden Age
might return—so the argument runs. The principal part of
this paper is an attempt to summarize the objective facts about
the relations between museums and the mainstream of an-
thropology in the past and at present, trying to strike a balance
between the bias of non-museum anthropologists who tend to
overlook the role of museums (especially in the past) and the
bias of museum anthropologists who tend to exaggerate the
importance of museums (especially in the present). Recogni-
tion of the objective situation is, I believe, a necessary pre-
requisite to policy decisions and to attempts at reformation. It
is especially necessary for museum anthropologists and mu-
seum administrators, whatever their wishes for the present and
hopes for the future, to admit the minuscule role and the low
prestige of museum work in present-day ethnology. Of course

52—Proc. Bro. Soc. Wasu., Vou. 82, 1969 (619)

620 Proceedings of the Biological Society of Washington

I believe that I am also correct in the value judgments I make
about the present situation and in the suggestions I present for
plans for the future. But I realize that these sections of the
paper may be considered controversial, and the reader should
evaluate the two parts of the paper separately.

If we adopt for a moment the usual, historically naive, ex
post facto outlook on the history of science, the beginnings of
anthropological collecting can be traced even before Aristotle
and Classical Greece. There is much archeological evidence
for the collecting of what would today be anthropological spec-
imens in prehistoric times—exotic objects and heirlooms have
been valued for almost as long as we have any evidence at all
on human culture. In more recent times, parallels to anthropo-
logical collections, and forerunners of them, can be seen in col-
lections of military trophies, in holy relics and the offerings of
the faithful kept in Greek and Roman temples and medieval
European churches, and in the powdered mummies, unicorn
horns, and other magico-medical items collected by early Eu-
ropean physicians and pharmacists. However, these collections
served motives and functions different from those of modern
museums. Collections of curiosities and archeological speci-
mens formed by the Chinese gentry and royalty in the 12th
century provide closer functional parallels to modern anthro-
pological collections (W. Trousdale, pers. communic., 9 Dec.
1968 ), but these are outside the historical tradition from which
modern anthropology and modern Western museums devel-
oped.

The real institutional beginnings of modern museums lie in
the Cabinets of Curiosities which came into vogue soon after
1500 A.D. (Murray 1904; Hodgen 1964: 114-23). The surviv-
ing catalogues and descriptions of these Cabinets show that
anthropological specimens formed a very important part of
them: many of the “artificial curiosities” (as opposed to the
“natural curiosities”) they contained would today be classi-
fied as anthropological, and the pieces in modern anthropolog-
ical collections which have the longest histories of continu-
ous preservation in collections are a few items which entered
Cabinets of Curiosities in the early 16th century, such as some

Natural history collection symposium 621

Mexican pieces sent back to Europe by Cortez after the con-
quest of Mexico in 1519 which survive in the Museum fiir V6lk-
erkunde in Vienna (Nowotny 1960).

Cabinets of Curiosities were important for the early develop-
ment of geology, biology, and archeology. The concept of
“archeological ages as technological stages” grew in large part
from the typological classifications of archeological artifacts in
Cabinets of Curiosities and in the first museums; the earlier rec-
ognition of the typological similarity between stone weapons
collected among contemporary North American Indians and
the “thunder stones” of European archeology provided an early
impetus to the notion of cultural evolution. But these Cabinets
were of practically no significance for the development of
ethnology, which grew instead out of written collections of
customs—compendia from travellers’ accounts and from classi-
cal literature of such things as religious customs and marriage
customs—a different kind of collecting, which began at about
the same time as Cabinets of Curiosities but independent of
them (Hodgen 1964: 123-206). There were no efforts to com-
pile systematic published accounts of the ethnological objects
in Cabinets of Curiosities, and very little attention was devoted
to developing logical classifications of these specimens (there
were of course published catalogues, and published collections
of such lists, but these show little or no effort to develop logical
or any other classifications of ethnological objects [Klemm
1837 contains a useful description and bibliography] ).

The beginnings of true anthropological collections in mu-
seums, the separation of these collections from other natural
historical and historical collections into distinct museum de-
partments of anthropology and into independent anthropolog-
ical museums, date from around 1840.! This was also the period

1 The precise dates usually given are often in fact rather arbitrary, for the older
museums evolved slowly by the amalgamation and subdivision of previous collections,
becoming distinct and public by a series of steps. However, the Ethnographical
Museum in Leningrad was established in 1836 (Troufanoff 1966: 232), the Na-
tional Museum of Ethnology, Leiden, dates itself from 1837 (Anonymous 1962: 3),
and the founding of the Ethnographical Collection of the National Museum of Den-
mark can be dated 1841 or 1849 (Birket-Smith 1968: 34-35). Frese (1960: 10)
gives a summary chart of the founding dates of European and North American an-
thropological museums (but the source of his data does not always distinguish the

founding dates of anthropology sections from those of the superordinate museum or
museum organization ).

622 Proceedings of the Biological Society of Washington

of the beginnings of modern anthropology with its emphasis on
the central importance of field research by the anthropologist
himself. The founding dates of the earliest professional societ-
ies of anthropologists fall into the same period (Société eth-
nologique de Paris, 1838; American Ethnological Society, 1842;
Ethnological Society of London, 1843).

What can be called the Museum Period of anthropology runs
from the 1840's to about 1590.7, During this time there was no
university training in anthropology, so anthropologists were
all people originally trained in other fields. Almost all anthro-
pological research was done by museum anthropologists, or by
amateurs, or by some other mavericks whose university teach-
ing responsibilities lay in other fields. Physical anthropology
was still largely a branch of human anatomy rather than a part
of anthropology, and most of its practitioners were associated
with medical schools. A nearly unique exception was the Bu-
reau of American Ethnology, which was founded in 1879 and
continued as a separate branch of the Smithsonian, administra-
tively independent from the U. S. National Museum (despite
the inauguration in 1883 of a Department of Anthropology in
the Museum). The staff of the B.A.E. conducted the most ex-
tensive and the most important anthropological research in the
United States during the last decade of the Museum Period and
the first decade or two of the ensuing period. The gathering of
museum collections during fieldwork, and studying them later
on in the museum, was however an important and respectable
part of anthropological research during this Museum Period.
The emphasis was on classification and typologies and geo-
graphical distributions. But museum collections were only mar-
ginally related to the development of theories of cultural evolu-
tion, which was the main focus of interest of anthropology during
this period. At the beginning, and in the prehistory of an-
thropology, typological studies of artifacts (both archeological
and ethnological) were important for the development of ev-

2 This periodization—Museum Period 1840-1890, Museum-University Period 1890—
1920, University Period 1920 to date—is developed from that implied by Collier
and Tschopik (1954). While it reflects primarily the United States situation, a
similar sequence obtains in other parts of the world. The second period probably
began two or three decades earlier in France and Germany, and lasted three or four
decades longer there and elsewhere in Europe.

Natural history collection symposium 623

olutionary theories—and also for the initial developments in
the now-discredited German “culture-historical” school. But
interest soon shifted to social evolution, and a good deal of the
most important anthropological work done during this period
had no relation to museum collections and could have been
conducted equally well if they had not existed at all: research
on kinship terminology, on the forms of marriage and the fam-
ily, on religion, has never depended at all on museum collec-
tions. Figure | shows that in the major German, British, and
American journals the proportion of ethnological articles which
made any reference to museum collections never rose above
20 percent during this period.

The next historical period of anthropology ran from about
1890 to about 1920. We can call this the Museum-University
Period. The formal teaching of anthropology in universities
began in the 1880's and 1890's in both England and the United
States, and in France, Germany, and the Netherlands rather
earlier (Quatrefages was appointed to a Chair of [Physical]
Anthropology at Paris in 1855 while Chairs in the Ecole d’An-
thropologie were inaugurated in 1875; Bastian was made Do-
zent fiir Ethnologie in Berlin in 1867; future administrators for
the Dutch East Indies received anthropological training from
1870). Still nearly all the jobs were in museums, most of the
teaching was done by anthropologists who also had museum
appointments, and museums supported most of the field work.
Museum collections remained important for research—in fact,
they became perhaps even more important, for the theoretical
developments of this period often used museum collections as
evidence, on such questions as the relative importance of dif-
fusion as opposed to independent invention, the relation be-
tween cultures and their natural environments, and in the ap-
plications of concepts from biology in developing the notions
of culture-areas and the age-area techniques of pseudo-histori-
cal reconstruction. The Bureau of American Ethnology con-
tinued to serve in effect as the research arm of Smithsonian
anthrepology; its collections were curated in the separate De-
partment of Anthropology of the U.S. National Museum, while
much of the publication and some of the fieldwork of the few
anthropologists in the Museum was supported by the B.A.E.

624 Proceedings of the Biological Society of Washington

In New York a somewhat similar relationship was worked out
by Franz Boas, the founder of academic anthropology in the
United States: between 1895 and 1905 he held a joint appoint-
ment in the American Museum of Natural History and at Co-
lumbia University, and used the museum as a base for his own
and his students’ fieldwork and its financing, from which mu-
seum collections resulted.

Yet the importance of museum collections for the anthropol-
ogy of this time should not be exaggerated. The chart (Fig. 1)
shows that in ethnology in the United States there was a steady
decline in their importance from a peak at 1900; the situation
in Great Britain and Germany is less clear, but here too such
collections were never the major focus of research. This was
the period of the rapid growth of fieldwork as the sine qua non
of ethnological research, and the collecting and study of ma-
terial objects played a relatively minor role in this fieldwork.
In archeology, too, such important developments of this period
as the application of the stratigraphic method were not derived
from work with museum collections as such.

In 1905 Boas resigned from the American Museum of Nat-
ural History in a conflict over the emphasis to be given re-
search; similar difficulties damaged anthropology in the Uni-
versity Museum in Philadelphia somewhat later. Darnell
(1968) has described these difficulties as conflicts between
the increasing professionalization of anthropology and_ the
growth of teaching departments with interests beyond ma-
terial culture, on the one hand, and the focus of museums (and
museum administrators and trustees) on objects, their collect-
ing, care, and exhibit, on the other hand. Beginning about
1920 we can speak of the University Period of anthropology,
which continues up to the present. With the gradual increase
in university teaching of anthropology, the balance shifted
until the majority of anthropologists was not employed in mu-
seums. The proportion of museum anthropologists has been
steadily declining, particularly rapidly during the last 20 years
with the really explosive growth of college and university en-
rollments in anthropology courses. Universities and founda-
tions took over the support of most fieldwork.

A measure of the relative importance for ethnology of mu-

Natural history collection symposium 625

seum collections, and of material objects whether or not in
museums, is the proportion of papers which touch on these
topics in the leading journals in the United States, Great Brit-
ain, France, and Germany (Fig. 1).* By this measure interest
in objects in American ethnology declined even more sharply
beginning in 1920 than it had before. In France, the decline
did not begin until the following decade—perhaps additional
evidence of the marginality of many aspects of French an-
thropology until after World War II which has recently been
noted by a French historian of the field (Mercier 1966: 104).
The corresponding decline in Great Britain did not begin until
1940; this is a surprising difference between British and Ameri-
can anthropology, which may indicate that the school of “social
anthropology” which came to dominate British anthropology
beginning about 1930 and soon had marked influences on
ethnclogy in the United States and elsewhere, was less an-
tagonistic to studies of material objects than is usually supposed
(e.g. by Hutton 1944, Collier 1962 )—or perhaps the dominance
was real but was inadequately represented by the editorial
practices of the journal examined. The German curves are of
very little significance for nearly 30 years following 1930; Ger-
man anthropology has only recently begun to recover from the
damage done to it by the Nazis.

This brings us to the present, where anthropology is in the
situation of having the responsibility for huge and irreplace-
able collections which represent a large investment over many
years of time, thought, care, and money, but seemingly have
very little importance for current anthropological research,
especially ethnological research. During the last 15 years,
North American anthropologists have published at least 10 pa-
pers deploring the situation of museum anthropology (actually
ethnology ) (Collier and Tschopik 1954; Shapiro 1958; Fenton
1960; Mason 1960; Collier 1962; Collier and Fenton 1965; Borh-
egyi 1965; Sturtevant 1966; Dockstader 1967; McFeat 1967);

3 For each country the joumal examined is the main vehicle of publication for
papers on ethnology without restrictions as to the geographical area or sub-topic
treated. The definition of ethnclogy applied in the counts is the one implied by an
exhaustive partition of anthropology into ethnology, archeology, linguistics, and physi-
cal anthropology. Abstracts, notices of meetings, book reviews, letters, and similar
brief communications, and papers on non-ethnological topics, were not counted.

626 Proceedings of the Biological Society of Washington

I870-79 1890-99 I910-I9 1950539" W950s59

eS %
N:27 liv 5 19 43 Ze oS at 42 OT
80b 80
7oL GERMANY moe 70
60 we 160
/ \
50. ; ee Fae
40, Ps Eee 40
30 ae | +30
20L ms 120
1OL 1O
GE %
is2 36 42 46 32 43 33 Zl a2 5|
60|. : 60
GREAT BRITAIN
50L _ eee as ie
a a 7
40} ao a ‘ 40
aia \
30/ oo Ae
20L 20
lob SS eee ee 10

FRANCE

Ne 10 44 44 43 53 7I 92 106 90
UNITED Fe
[ STATES

~
SN

1880-89 1900-09 1920-29 1940-49 1960-67

Ficure |. Interest in Material Culture and Museum Collections in
Ethnology. Dashed lines show the percentage of all papers on ethnology
which are concerned (at least in part) with material culture; solid lines
show the percentage of all papers on ethnology which are based (at least
in part) on museum collections. Sources: American Anthropologist

(1888-1967); PEthnographie (1913-1965) and THomme (1964, 1967):

Natural history collection symposium 627

the most extensive treatment of this and related problems is a
monograph by a Dutch anthropologist (Frese 1960). I know
of no British, French, Scandinavian, or German papers which
parallel these (although they may exist), but conversations
with museum anthropologists from these countries over the
last year or two have convinced me that the situation is not
very different in Europe.

Although it is customary to write about “anthropology” and
museums, in fact some distinctions between the sub-fields of
anthropology must. be drawn before a sensible answer can be
given to the question posed at the head of this paper. Anthro-
pology is quite sharply divided into four sub-fields, and one of
the most marked differences between them is the use they make
of museum collections. These four sub-divisions are linguistics,
physical anthropology, archeology, and ethnology.!

The relation between linguistics, the scientific study of lan-
guage, and the usual museum anthropological specimens, is nil.
This is true of anthropological linguistics, which is based on the
field study of languages which are still spoken and to a lesser
extent on written records of them made in relatively recent
times. The U.S. National Museum is perhaps unique among
museums in including in its collections extensive linguistic
archival materials useful for anthropological linguists. Those
linguists who study extinct languages known only or largely
through documents recovered archeologically—for example

4 This represents more or less standard American usage, except that (for good
reasons) I prefer the somewhat old-fashioned and museum-oriented label “‘ethnology”’
for what is often now called ‘“‘cultural anthropology” or “social and cultural anthro-
pology.” In Europe these four fields (and folklore) are less often viewed as com-
ponents of a single larger discipline. Tendencies in Europe towards integrating the
fields and in America towards incorporating into anthropology studies of Euro-
American cultures have as yet had little effect on the organization of museums,
whose buildings, collections, and bureaucracies cause them to lag behind universities
in the reorganization, amalgamation, and subdivision of traditional departments.

>

<

Journal of the (Royal) Anthropological Institute (1872-1964) and Man
N.S. (1967); Zeitschrift fiir Ethnologie (1871-1967 ). Every third volume
of each journal was scored (with some adjustments for France during
W.W. I and Germany during W.W. II); these scores were then lumped
by decade. N = number of papers on all ethnological topics in that dec-
ade’s sample.

628 Proceedings of the Biological Society of Washington

Mesopotamian clay tablets or Egyptian inscriptions or papyri
are not anthropological linguists and the museum specimens
they study are only rarely kept in anthropological museum col-
lections.

Physical anthropology deals with human biology. Thirty and
more years ago, osteometry based on museum collections was a
major interest of the field (although anthropometry of the liv-
ing was also important). In recent years sub-specialties such as
human genetics and primate ethology which have little or
nothing to do with museum collections have been growing, and
classical osteometry and anthropometry have nearly dis-
appeared. Research on human paleontology, paleodemogra-
phy, and paleopathology still depends on skeletal material, but
the older museum collections are often of little value ( especially
for demographic studies ) because they rarely constitute proper
samples of the ancient populations. Present research concen-
trates on newly excavated materials, and the new necessity to
keep even fragmentary specimens sometimes poses storage and
cataloguing problems.

An indication of the relation of museum collections to re-
search is the proportion of new accessions which come in
without specific data on their sources. Practically no such
specimens are now accepted into the physical anthropology
collections in the U. S. National Museum; bones not accom-
panied by precise information as to their spatial and temporal
provenience are not worth accepting and preserving, because
they cannot be used for research.

Specialists in physical anthropology are a small minority of
the total number of anthropologists, and very few museums
maintain collections in this area. It is, however, becoming dif-
ficult to find properly qualified curators for these collections
since the research of most physical anthropologists no longer

depends on museum specimens.

Archeology, which is the study of fossil cultures, of cultural
evidence recovered largely through excavations, is the part of
anthropology for which museum collections are most important.
The whole subject rests directly on the study of material ob-
jects and material remains, used as evidence for deductions
regarding the human past. Of course the purpose of research is

Natural history collection symposium 629

not the simple amassing of museum specimens—an_ activity
which archeologists call “pot hunting” and consider to be mere
vandalism. Rather, advances come through new fieldwork, new
methods of observing, recording, and interpreting, and the pub-
lication of these results. However, most archeologists consider
that a major part of their responsibilities for documenting their
results consists in providing a properly catalogued museum col-
lection, because publication alone does not provide adequate
data for future research, which must continually check back
with previously excavated specimens in order to set the new
work into context and in order to reinterpret the old results in
terms of new typologies and new descriptive techniques. Ar-
cheology thus has an important “taxonomic” base in museum
collections, much like some of the natural sciences.

As with physical anthropology, undocumented specimens are
normally not accepted into museum collections. In recent years
well over 90 percent of the archeological specimens added to
the U.S. National Museum collections have come from excava-
tions by professional archeologists. Furthermore, archeologists
have little hesitation in deciding what parts of their field col-
lections should be kept in the museum collections and what
parts can be discarded after they have been recorded. Hind-
sight sometimes shows that mistakes have been made, but the
central position of material objects in the research means that
at a given period there is good agreement on what must be
kept for documentation.

Even though most current research depends on new field
studies, there remain many important museum collections re-
sulting from older excavations which have never been ade-
quately studied. The occasional archeologist who analyzes and
publishes these old collections is not felt by his peers to be
wasting his time, and such studies can be expected to increase
with the rapid destruction of archeological sites in many parts
of the world in the construction of dams, highways, and indus-
trial plants, the expansion of cities, and the increasing use of
earth-moving machinery in agriculture. Much of Classical
archeology already depends on the study of existing museum
specimens, often with inadequate contextual data (this is one
of the respects in which this field is peripheral to, or outside,

630 Proceedings of the Biological Society of Washington

anthropological archeology ). On the other hand archeologists
working in some parts of the world are forced to do without
museum collections because they are prohibited from export-
ing their excavated materials while local museums are still un-
able to preserve them for future research.

Ethnology, the fourth sub-field of anthropology, is the study
of living cultures, especially by means of the sort of fieldwork
known as ethnography, which requires participant observation
(extended periods of face-to-face relations with members of
the society being studied, observations and interviews con-
ducted on the spot by the ethnologist himself). A minor strand
in ethnology makes use of contemporary written documents
about now-extinct societies or the past stages of existing socie-
ties, but this “ethnohistory” depends heavily on methods de-
veloped by ethnographic fieldwork.

Ethnology is today the central field of anthropology, the one
which holds together the four sub-fields. Anthropological lin-
guistics, archeology, and physical anthropology are parts of an-
thropology largely by virtue of their interrelations with ethnol-
ogy, and particularly because of the central position held by
the (ethnological) concept of culture in definitions of the cov-
erage and the methodological and theoretical emphases of the
non-ethnological sub-fields. There are some kinds of linguis-
tics, archeology (or prehistory ), and human biology which are
non-anthropological in terms of the methods, interests, train-
ing, and professional self-identification of their practitioners,
while there are no professional ethnologists who are not an-
thropologists in this sense. This formulation—which is probably
acceptable to most non-ethnologist anthropologists, at least in
North America—does not deny the fact that linguistics, archeol-
ogy, and physical anthropology have varied relations between
each other and with disciplines outside anthropology. For ex-

ample, archeology is more closely dependent on several of the
natural and physical sciences than is ethnology, and in turn can
contribute to their historical aspects in ways that ethnology
cannot. It is also true that many of the interests and methods
of ethnology depend on contributions from the other fields of
anthropology, and from other disciplines such as psychology,
sociology, economics, and history. But anthropology remains a

Natural history collection symposium 631

single subject, with sub-divisions. Some observers believe that
it will not (and sometimes that it should not) remain so, that
increasing specialization will lead to fragmentation. But this
specialization often overlaps sub-field boundaries, so that the
discipline may well become a network rather than a rigid set
of four pigeonholes. I believe that the sub-fields will (and
should) continue to offer more to each other than to outside
disciplines. If museums need anthropology, they must include
ethnology.

But ethnology is the anthropological sub-field which has the
most ambiguous relation to museum collections. Ethnologists
study culture, and they often boast that, in contrast to practi-
tioners of the other social sciences and humanities, they study
both all cultures and all aspects of culture. A classification of
the aspects of culture useful for present purposes is a common
one which distinguishes three major classes: material culture,
social culture, and mental culture. To characterize these
roughly, material culture is concrete artifacts or manufactures,
social culture is behavior, and mental culture is ideas, knowl-
edge, and beliefs.®° Only material culture can be represented in
museum collections, and it is perfectly possible—indeed it is
usual—to study social and mental culture without paying any
attention to material culture, to artifacts, and therefore to mu-
seum collections. Material culture studies themselves are of
course not limited to work with museum collections, for the
contexts of the objects in the social and cognitive systems of
their makers and users is a primary interest.

As with the other sciences represented in natural history mu-
seums, collections are relevant to only some kinds of anthropol-
ogy and often not to those areas in “the forefront of research”
(cf. Crompton 1968). But there is a significant difference: for
the core area of anthropology, “systematics” and “basic de-
scriptions” based on or documented by museum collections are

5 See Osgood 1951 for these categories, defined on a somewhat different basis.
The definition which I prefer for both theoretical and methodological reasons puts
the locus of ‘‘culture”’ in the minds of its bearers, which makes the term “mental cul-
ture” redundant and requires rewording of the labels for the material and social re-
sults of culture: perhaps ‘‘cultural materials” (i.e. artifacts) and ‘‘cultural behavior.”
If artifacts are thus viewed as reflections of culture rather than part of culture, they
are of no less value as documents or evidence on a major aspect of culture, on the
varieties of specifically human cognition and behavior,

632 Proceedings of the Biological Society of Washington

not now and have never been fundamental in any sense to other
research. Artifacts and museum collections of them play no role
as ethnological “standards” or “vouchers”; the units of ethnolog-
ical study are bounded, identified, and classified without re-
gard for museum collections. This would not be important for
museum anthropology if ethnologists were really equally inter-
ested in ali aspects of culture. But as has already been in-
dicated, this is not the case. From the beginning, research on
material culture has been less important in ethnology than
research on social and mental culture.

In 1967 the three major general anthropological journals in
the United States, England, and France published 65 papers
on ethnological topics. Of these, only five dealt with material
culture; among even these, three were based on field observa-
tions and made no reference to museum collections. The over-
whelming majority—60 to 63 out of 65—could have been writ-
ten if there were no anthropological museum collections at all.
Even the research of most museum ethnologists does not in-
volve material culture or museum specimens. Most modern eth-
nologists have never studied museum specimens, have never
collected for a museum, have never even been in a museum
storage area. Yet I suppose at least 90 percent of museum eth-
nological specimens have never been studied.

In a few decades, anthropologists will surely look back on
the present time as the last period when it was possible to col-
lect hand-made traditional artifacts, and to document their pro-
duction, local terminology, and uses by field studies, before
they were completely replaced by mass-produced manu-
factured goods of the “international style.” Nearly every eth-
nographer could collect now; hardly anyone does. No anthropo-
logical museum seems able and willing to provide funds to en-
courage collecting by the hundreds of ethnographic field
researchers now at work. The budgets of most museum anthro-
pology departments do not regularly include sufficient funds to
purchase even the useful collections which are offered. When
funds are available, high prices tend to go for showy pieces
without documentation bought on the art market. If items col-
lected by a trained ethnographer with proper scientific docu-
mentation can be bought, the price paid normally covers only

Natural history collection symposium 633

the actual costs of purchasing, packing, and shipping the speci-
mens. Yet there are many ethnographers (especially outside
the United States) who lack sufficient funds to support their
own fieldwork and who would readily devote some extra time
and attention to making a properly documented collection if
they were offered a reasonable mark-up over their out-of-
pocket expenses, which could be used to help pay for their other
work.

As recently as ten years ago, an ethnologist on the British
Museum staff wrote that in the United Kingdom, “collecting in
the field is rarely possible for most museum officials in charge
of ethnographical collections” (Cranstone 1958: 7), and the
situation has changed little since then. In the United States
and a few other countries funds are not so short and the poli-
cies of large museums regarding fieldwork by their staffs are
not so restrictive. Yet over the last four years, nearly two-thirds
of the specimens added to the ethnological collections in the
U.S. National Museum were not collected by ethnologists, but
were collected under non-scientific conditions by untrained
people and hence lack essential documentation as to proveni-
ence, age, functions, and so forth. Of course non-anthropologists
can collect materials which are scientifically useful. However,
a set of directions and suggestions on how to make an adequate
field collection of ethnographic specimens which the U. S.
National Museum published in 1967 was the first such guide
published in English since 1902; the last one in French is dated
1931 and the last in German, 1914 (Sturtevant 1967; Holmes
and Mason 1902; Musée dEthnographie 1931; Ankermann
1914).

The relative unimportance of collections is demonstrated by
the growing tendency to separate them from the associated
scientific staff, public exhibits, museum administrative space,
and classrooms. The more convenient centrally located space
is repeatedly being found to be too valuable to use for storing
specimens. But if the specimens were really significant for re-
search, it would be as inconceivable as it is for research librar-
ies to locate them several miles away from the researchers
(usually without plans for a regular service to transport people
and objects between the two locations). What is objectionable

634 Proceedings of the Biological Society of Washington

is not the separation of the collections from the exhibits, but
storing the specimens miles away from the associated records
and the scientific staff.°

As Crompton (1968), Washburn (1967, 1968), and others
have pointed out, when research on collections is infrequent
and of low prestige museums naturally seek other justifications
for existence—popular exhibits, general education—and the
staff members tend to become administrators, showmen and
public relations experts, and museologists, rather than subject-
matter specialists. The results for research on the collections
and even for their preservation are obviously disastrous; that
this is not hypothetical can be seen from the history of many
museums (see, e.g., the cases described in Whitehill 1967 ). An-
thropological collections are even more liable than some others
to suffer, for many kinds of anthropological specimens require
constant attention to prevent deterioration, many are of high
value on the art market, and research on them is at a particu-
larly low ebb. Some recent examples of the results are perti-
nent: a naturalist in charge of a museum overrides his anthropol-
ogist curators and authorizes the loan of important ethnological
specimens for decorating politicians’ offices; an ethnologist
museum director sells unique ethnographic specimens cata-
logued in his museum, both at the public sales desk at his insti-
tution’s front door and through profit-making dealers in “primi-
tive art”; one archeologist museum director trades important
well-documented early ethnographic specimens from his mu-
seum to a private individual in exchange for an easily dupli-
cated collection of non-excavated archeological sherds; another
archeologist in charge of a museum orders each of his curators
to select specimens for sale at a private auction to his socialite
“friends of the museum”; one major anthropological museum
charges visiting researchers $50 to open an exhibit case in order
that displayed specimens may be studied; an ethnologist chair-
man of a department in another museum suggests that a quali-
fied visiting student prepare the first thorough descriptive cat-

6 Such plans for removing the anthropological collections are in various stages of
completion at least in the British Museum and the Homiman Museum in London,
the Peabody Museum at Harvard, and the U. S. National Museum. The Museum of
the American Indian in New York has operated with such a separation for many
years.

Natural history collection symposium 635

alogue of one of the most important collections under his care,
and then refuses to allow the student to complete the catalogue
by including those pieces in the collection which have been
solidly built into modern exhibits on the grounds that it is too
much trouble to remove them for study. As one of the small
group of research users of ethnographic collections, my own
experiences on study visits to some 15 of the 20 or so largest
and most important general ethnographic collections in the
world are significant. Two of these museums flatly refuse to al-
low serious researchers to photograph their specimens; most
have no special facilities for visitors to use for photography, and
many have not even any space where a visitor can arrange
items to photograph even though he has brought all his own
equipment; none, in my experience, has convenient locations
for studying the specimens in or immediately adjoining all stor-
age areas; most find it difficult—and some impossible—to re-
move exhibited items for study (but all try to put their most
important specimens on exhibit, often with catalogue numbers
hidden ); usually some 10 to 20 percent of the specimens a visitor
selects for study from the catalogue descriptions cannot be lo-
cated (and in a recent visit to a national museum of anthropol-
ogy in Europe, 83 percent of the specimens I identified in the
working catalogue could not be found); always a visitor can-
not help but feel that he is imposing on the inadequate profes-
sional and supporting staff—a visitor interested in serious re-
search on the collections is so unusual that he is bound to
disrupt the museum routines. The usual state of the storage and
the catalogues and other records has to be seen to be believed;
one seriously wonders whether present collections will survive
any better than have the pitiful remnants of 17th and 18th
century collections (cf. Washburn 1968 ).

But let me switch hats to my role as curator. An ethnologist

with curatorial responsibilities, while recognizing these dis-
graceful conditions, must also consider the allocation of his
own time and energies. What should be done to improve and
preserve the collections is obvious; but the results of his work
would be seen and appreciated by a very small proportion of
his colleagues, and given the severe limitations in funds, per-
sonnel, and space all museums suffer from, it would be a dif-

636 Proceedings of the Biological Society of Washington

ficult struggle to get even a small part of the help so obviously
needed to do a proper job. The criteria by which his professional
standing is evaluated both by his anthropological peers and by
the museum authorities who pay and promote him have almost
nothing to do with the state of the collections under his care.
Curators with any ambition and regard for their own potentiali-
ties quickly and repeatedly decide to devote themselves to the
research and publication which will advance anthropology
(and their own careers ) right now rather than in some distant
future. Such ethnologists are “square pegs in round holes” or
“in the wrong pew —to quote the common opinion of museum
archeologists and of the few really good and productive mu-
seum ethnologists who do focus their research on the collec-
tions under their care. But there are nowhere near enough
good round pegs to fill the holes in museums. The alternative to
supporting square pegs is to hire museum ethnologists who are
not in the mainstream of ethnology, which further degrades the
attractiveness of museums for active anthropologists of what-
ever specialty. There are a few such people now in museums;
among them are some of the better curators, but also some of
the worst: lacking peers, they are less constrained by outside
judgments of their actions and easily fall into autocracy, isola-
tion, high-handed treatment of research visitors, and disposal
of scientifically vital collections through sale or exchange to
individual collectors and dealers and to other (especially art )
museums. The administrative structure of many independent
and some university museums only supports these tendencies,
tor boards of directors and boards of supervisors tend to con-
sist of financiers, businessmen, politicians, and others who are
interested in the financial status of the organization and in its
reception by the general public, but who cannot and do not
exercise any informed scientific supervision over a director
gone berserk.

What can be done? It is a problem for museum anthropology
as a whole, not just for museum ethnology. Although collections
are central to the research of archeologists and some physical
anthropologists, but only to a very small minority of ethnolo-
gists, the answer is not to separate out the archeologists and
physical anthropologists and their collections. Not only would

Natural history collection symposium 637

this be disastrous for museum ethnology, but it would be dele-
terious for museum archeology and physical anthropology, for
anthropology is fundamentally a single field and few anthro-
pologically-oriented archeologists and physical anthropologists
would stay in fragmented departments where they would be
peripheral to the centers of unified anthropological research
and teaching.

The best hope is for the increase of the quantity, quality,
and prestige of ethnological research based on museum collec-
tions. Broad justifications for the importance of ethnological re-
search on material culture (which in turn will require attention
to artifacts in museums ) are not difficult to formulate:

1. Man is preeminently the tool-using animal, so that an un-
derstanding of his physical and cultural evolution and his rela-
tion to the non-human environment requires knowledge of his
adaptive use of materials in its full cultural variety in historic as
well as prehistoric times.

2. Ethnology is not fulfilling its mandate when it neglects
material culture in favor of social and mental culture. In many
respects the material basis clearly underlies, limits, and deter-
mines other aspects of human social life. It is particularly sur-
prising that the technological aspects of our own and other cul-
tures are not more studied by anthropologist members of a
society so dominated and harassed by technological advances
and technological problems. If anthropologists do not fill this
gap, it will be filled by others who lack some of the special ad-
vantages of an anthropological training and outlook, in partic-
ular the emphasis on functionalism which leads to studies of
the integration of artifacts with non-material aspects of cul-
Lune:

3. Artifacts, and especially dated artifacts in museum col-
lections, provide essential evidence for the history of cultures.
Ethnological artifacts are an important link between the socie-
ties whose remains are recovered in the more recent parts of
archeological sequences, and their historical successors. Further-
more, archeologists depend heavily on ethnological analogies

7 The last two points were emphasized for me in conversations respectively with

P. J. C. Dark and J. C. Ewers.

635 Proceedings of the Biological Society of Washington

for understanding the functions and contexts of the fragmen-
tary artifacts on which they must base their paleo-ethnography
and prehistoriography. For ethnohistorians museum collections
are crucial historical documents whose potentials are only
beginning to be appreciated (cf. Fenton 1967 ).

4. In non-literate societies only artifacts provide models and
evidence of the past apart from those “stored in human mem-
ory (and subject to the vagaries of human memory); this
surely has important consequences for the members of those
societies (Goody and Watt 1968: 29), as it certainly does for
the evidential value of artifacts for both contemporary and
subsequent outside observers. Both informants’ and recorders’
biases are less significant here than with either oral or written
testimony. The artifacts stored in museums provide a vast body
of quite direct cultural evidence which should be analyzed
and re-analyzed.

But general statements such as these on the importance of
material objects for human life, and on how unjustifiably mu-
seum collections of them are being neglected, are not going to
convince students nor shift the research interests of established
professionals. When the statements come from a museum an-
thropologist they sound like petty and self-serving complaints
which are easily taken as attempts to denigrate the real ac-
complishments and importance of other more active lines of
current research. What causes shifts in research emphases is
the discovery of quite specific problems and methods that are
attractive because they promise advances clearly related to
other important interests of the discipline. If such problems
and methods can be worked out from studying museum col-
lections, this in turn will raise the prestige of more pedestrian
research done on the same media. Attention should therefore
be devoted not just to urging more research on artifacts, but to
improving the methods of research on museum collections and
particularly to adapting interesting developments from other,
more prestigious and more advanced fields. Such applications
are more likely to be made by the “square pegs” with other
interests whose employment puts them into proximity to the
collections, than they are by “round pegs” attracted to museums
by the traditional kinds of research on ethnological collections.

Natural history collection symposium 639

In fact, there are already indications from several different
directions of a revivification of ethnological research on ma-
terial culture. While this is not the place to go into details, an
enumeration of some of these tendencies (or potential tenden-
cies ) helps to justify optimism about the future of anthropologi-
cal museum collections. From archeology may be mentioned
the application of attribute analysis to ethnographic specimens
as well as archeological ones, and the increasing importance of
detailed and specific ethnographic analogies in archeological
interpretations. From other interests of ethnology (and _lin-
guistics ) come: recognition of the advantages of concrete arti-
facts as the basis for componential analysis and for other ap-
plications of etic/emic or ethnoscientific methods, generative
analysis, semiology, and other semantic approaches; the in-
volvement of art and artifacts in studies of symbolic classifica-
tion; an increased interest in field studies of non-Western art,
from various points of view (partly influenced by lessening
ethnocentrism in Western art appreciation and art history );
the use of specimens, especially dated ones, as historical docu-
ments on both non-literate and literate cultures; and the rec-
ognition of the utility of artifacts in museum collections for
the critical assessment of ethnographic illustrations both as
ethnological documents and as part of the history of Western
art. These trends may be summed up as an increasing atten-
tion to classification, semantics, and symbolism—in general, the
rise of a variety of structuralist methods—and in diachronic
studies more inclusive definition of the kinds of “documents”
which are relevant.

It is not only developments in anthropological theory and
method that encourage confidence in the wider recognition of
ethnological museum collections as the important resources
they indeed are. In France, at least, there is already an ob-
vious increase in student interest in material culture and mu-
seum collections: one of the demands of the protesting students
of May and June 1968 was for access to museum collections and
introduction to their study (Hélene Balfet, pers. communic., 15
Feb. 1969). The combination of the increasing difficulty of
access to foreign areas for fieldwork, the very rapid Westerni-
zation of technology everywhere, and the explosive increase in

640 Proceedings of the Biological Society of Washington

the number of anthropologists who must publish or perish, will
almost certainly also lead to more research on ethnological mu-
seum collections.

Meanwhile there are several organizational modifications
which can improve museums as research environments for an-
thropologists and help to save their collections for the time
when they will be vital for anthropology.

Museum specimens are unique cultural and historical docu-
ments; we must find out what and where this evidence is.
There are about 200,000 ethnological specimens in the U.S.
National Museum, somewhat over one and one-half million in
all United States museums, and perhaps four and one-half mil-
lion in all museums of the world.’ A pilot study at the Uni-
versity of Oklahoma has developed procedures for preparing
an inventory of all of these, which would incorporate most of
the errors in existing museum catalogues (for example, the
U. S. National Museum must have several hundred, perhaps
several thousand, specimens catalogued as “locality unknown,
probably North America” or some equivalent of this) but
would provide the basis for later correction and amplification.
This study indicates that it would require about 140 man-years
to prepare an “index ethnographicum” or “union catalogue” for
the United States alone, at a cost of approximately 50 cents per
specimen for preparing and key-punching the inventory sheets
any computer operations will add to this cost figure ( Ricciar-
delli 1967b, 1967c). Somewhat over half the specimens in the
United States are in the five largest museums, which should
surely be left to do their own indexing; they have or can get
the needed skilled staff, and this will make partial completion
of the project less than half as expensive and nearly as useful as
full completion, for anyone will know that he must search these
major museums for relevant specimens whereas without an in-
ventory he will miss most of the others which are widely scat-
tered in smaller museums. As soon as possible these large mu-
seums should modify their present cataloguing systems to make

8 These figures are based, respectively, on (1) a careful count of a stratified sam-
ple of the specimens described in the USNM catalogue cards, conducted by the
author and Gordon D. Gibson in 1965 and 1966; (2) the North American estimate
made by Ricciardelli (1967a) from several lines of evidence carefully considered; (3)
my own extrapolation from the latter, which is merely an informed guess.

Natural history collection symposium 641

them compatible with the projected continent-wide computer-
ized index, so that future accessions can be fed into the system
immediately, before the index is extended backward to include
the older materials. Similar schemes are being considered at
least in the United Kingdom and France; there is reason to
hope that all will be compatible. As Jean Cuisenier has pointed
out (pers. communic., 30 Dec. 1968), the use of computers is
spreading so rapidly that the modern student generation takes
them for granted; our museum collections are in danger of be-
coming useless if young scholars are not able to use computers
to retrieve information on them.

Most research on ethnological collections depends heavily on
the minority of specimens which have some documentation, at
least dating, and the older collections of this sort are particu-
larly valuable. So a committee of the International Council
of Museums (ICOM) and J. C. Ewers with the Committee on
Anthropological Research in Museums of the American Anthro-
pological Association are both considering another type of in-
ventory to compile location lists for older dated specimens
without waiting for these to appear in the full inventories of all
museum ethnological holdings.

The problem of the conflict between curatorial and research
duties is perhaps even more acute in ethnology than in other
museum fields, because of the wider gap between the usual re-
search interests of present and prospective curators and their
housekeeping responsibilities. Complete separation of research
and curatorial staff is risky: in many if not most museums the
collections and the necessity for exhibiting and caring for them
provide the front which justifies the museum budget; if it is
made to appear that research and curation are completely dis-
tinct, research becomes more vulnerable to budget cuts; but it
is well known from much experience that collections without
associated research staff cannot long survive. On the other
hand, giving the research staff full curatorial duties has the
untoward consequences for both the collections and the re-
search which we have already outlined. One solution is to de-
velop further the practice already existing in most large mu-
seums, where the scientific staff supervises a “supporting staff,”
paid less and with lower academic credentials, which does most

642 Proceedings of the Biological Society of Washington

of the actual curatorial labor. But it is difficult to locate, train,
and keep adequately skilled people for such clearly second-
class jobs. The status and responsibilities of these positions
could be raised by lifting the career ceiling on them and assign-
ing to their upper ranks some such title as “Curator of Collec-
tions,” with truly commensurate responsibilities. A few mu-
seums already do this, and the practice should be extended. We
need the museum equivalent of Librarians and Archivists. Pro-
fessionalization of this sort does carry the dangers that Wash-
burn (1967) has pointed out. The scientific staff—subject-
matter specialists—must maintain scientific guidance over
collections policy, and museum tables of organization should be
planned with this in mind. It may be anachronistic in this so-
ciety, but an effort must be made to emphasize apprenticeship
training rather than preparation in some academic museology.
Certainly the knowledge and experience needed to curate mu-
seum collections is more specialized, more different as between
the collections of different sciences, than is the case for col-
lections of books or manuscripts. An anthropological museolo-
gist, an entomological museologist, and an art-gallery museolo-
gist could not come from a similar background of academic and
practical experience.

Finally, some important modifications of the museum concept
are needed at least by anthropology. For one thing, anthropol-
ogy does not belong in a natural history museum. In fact, the
United States is behind the rest of the world in this respect: ex-
cept in North America, Australia, and New Zealand, nearly all
important anthropological collections are either housed in in-
dependent museums of anthropology or of man, or they are
joined with collections of history, folklore, prehistory, and Clas-
sical archeology, while natural history collections are separately
housed (Frese 1960: 15-32). A justification for the separation
which is of particular force for the modern world is that given
by the Director of the National Museum of Anthropology of
Mexico in describing its origins in 1910: “Until that year the
museum had remained one of “Natural History.” But at that
time all the natural history collections were removed to another
museum thus abandoning, I hope forever, the placing of indige-
nous cultures in the same building as animals, which gives visi-

Natural history collection symposium 643

tors inaccurate ideas about native peoples and their cultures”
(Bernal 1966: 132). Another reason for removing anthropology
from natural history museums is the quite different character
of the collections, which are more like those of history and art
in their unique qualities as historical documents and in the
problems of acquisition and protection, and which are related
to ongoing research in quite a different manner. Some anthro-
pologists (especially some archeologists) now in natural his-
tory museums point to the advantages of a close association
with the natural scientists with whom they find many areas of
scientific collaboration, especially with the rise of an ecological
approach to human cultures. But there are equally strong rea-
sons, from some other areas of anthropological interest, for
urging the benefits of a closer association with the historians,
art historians, and technologists who are found in other kinds of
museums. Another advantage of a separate Museum of An-
thropology or Museum of Man is that it is easier to broaden its
mandate for collecting and curating so that it will include all
the sorts of physical objects on which anthropological research
is based. The Musée de [Homme and the Musée des Arts et
Traditions Populaires in Paris, and the Department of Anthro-
pology and the Center for the Study of Man (now planning a
new Museum of Man) of the Smithsonian, and probably a few
other museums, already define their museum function as es-
sentially that of archiving: the usual museum collections of
artifacts and skeletal materials, and in addition still and cin-
ema photographs, drawings and paintings, sound recordings,
anthropological manuscripts, and books. Many of these addi-
tional materials are at least as crucial for future research as are
specimens and yet are not being systematically archived by any
other institutions; the physical and administrative museum
structure is more suitable for this task than is that of any univer-
sity department.

With new museums comes the rare opportunity for a major
advance in anthropological exhibit techniques. Any museum
anthropologist will recognize the advances associated with the
inauguration, in order, of the Pitt Rivers Museum in Oxford,
the Natural History Building of the U. S. National Museum,
the American Museum of Natural History, the Musée de

644 Proceedings of the Biological Society of Washington

Homme, and lastly, the Milwaukee Public Museum in 1963-71
and, in 1964, the new building of the National Museum of An-
thropology in Mexico City. It is past time for a radical new ap-
proach. Borhegyi has recently well described the problem:

Through [museum] exhibits, million of people can be ex-

posed to the inherent dangers of nationalism, ethnocen-

trism, and racial and religious prejudices. Yet museum ex-
hibits in general, and natural history museums in particular,
instead of stirring the imagination of visitors, tend to
perpetuate the visitors’ stereotypes of “savages” and

“quaint primitive” cultures. The anthropology exhibits

keep on cultivating the romanticism of the visitor by

showing exotic “tribal” peoples in “peculiar” attires, amidst
prettily staged sentimental settings, or appeal to his sense
of the macabre by the inevitable showing of mummies,
skeletons, and shrunken heads. . . . Museum anthropolo-
gists continue to be primarily object and tribal rather than
subject or concept oriented in their exhibits, and most of
them rightfully deserve the title of “keepers” . . . rather
than “doers.” (Borhegyi 1969 )

Perhaps three new approaches to exhibits would be particularly

effective in anew Museum of Anthropology:

1. Exhibits should catch up with the principles of modern
anthropology, rather than continuing simply to illustrate the
“culture areas” elaborated for museum exhibits over 60 years
ago. In particular the relevance of anthropological knowledge
to some of the difficulties of the modern world should be
stressed.

2. Some exhibits, perhaps changing ones, should illustrate
current research, especially that being conducted by anthropol-
gists on the museum staff.

3. Anthropology, as the only social science well established
in museums, seems the ideal field to study the educational ef-
fectiveness of various exhibit techniques, to conduct research
on visitor reactions. I am by no means an expert on the topic,
but I have the impression that this is an underdeveloped
research area. The rapid specialization and technological im-
provement of exhibit techniques seems to have occurred with-
out reference to studies of what visitors actually prefer or bene-

Natural history collection symposium 645

fit from (and these last may not be the same). A recent wide-
ranging bibliography of museum visitor surveys lists only 124
titles, published and semi-published, nearly all of them very
brief papers, on studies in all sorts of museums between 1597
and 1966, and very few of these report anything approaching
sophisticated controlled experiments (Borhegyi and Hanson
1968a; cf. Washburn 1961; Borhegyi and Hanson 1968b ).

Although anthropological museum exhibits certainly need
improvement, there is a real danger that attention to exhibits
will intrude on the time and support for curators’ research.
Certainly no exhibit program should be conducted without
both the technicians to do the actual work, and funds to hire
outside experts on a short-term basis to help plan the scientific
aspects of the exhibits. Down the exhibits road lie the mu-
seums feared by research-oriented curators, where emphasis on
exhibits, popular education, visitor attendance, advertising, and
income-producing museum shops erodes support for scientific
research, drives scholars off the staff, and runs a grave risk of
destroying the collections and turning the museum into a mere
entertaining sideshow.

Crompton has recently urged that “it is time .. . that we rec-
ognized that the functions of maintaining collections, design-
ing exhibitions and running sophisticated research programs
cannot be carried out by a single person. It must also be rec-
ognized that successful scientific research is usually coupled
with stimulation provided either by fellow workers or students
or teaching or all three. Unless natural history museums are
prepared to recognize this, it will not be possible for them to
create strong scientific programs.” He also outlines the manner
in which a successful university museum may avoid many of
the problems of ensuring active research by its curators, by
integrating the museum administratively with the teaching de-
partments (in reality, subordinating the museum to the depart-
ments ) (Crompton 1968). A non-university museum must in-
vent the equivalent of teaching departments. Opportunities
must be provided for curators to take leave to teach in univer-
sities, and fellowships and facilities must be offered to attract
students and university faculty members to museums—and not
only for research and teaching related to the museum collec-

646 Proceedings of the Biological Society of Washington

tions. Museums have some advantages over universities as
bases for anthropological research; among these is the freedom
from the academic schedule which allows extended fieldwork at
any time of the year. Particularly in ethnology it is customary
for the most intensive and important fieldwork to be done early
in the scholar’s career, often just before he receives his Ph.D.
In the usual situation he must then postpone publishing the full
results; by the time his teaching duties become less time-con-
suming and he can get leave from his university, he has family
and other responsibilities which prevent another lengthy period
of isolation for fieldwork. The same is true, to a somewhat
lesser extent, for the other sub-fields of anthropology. Itis becom-
ing ever more clear that advances in ethnology depend on
advances in ethnography; yet ethnography suffers from the
structure of academic careers. Museum-based research, both
fieldwork and publication, for younger anthropologists is a solu-
tion. If after a few years they move to university teaching posi-
tions, the museum and the science have gained by supporting
them during their most productive research years, and the
university has gained by acquiring teachers who are already ex-
perienced and productive research workers.

As Fenton (1960) has suggested, a redefinition of anthropo-
logical museums in terms of the Alexandrian museum as a com-
munity of scholars and students would be a large step forward.
Collections will be increasingly important, and there are se-
rious problems in preserving them and in taking advantage of
the short time remaining in which we will be able to use field-
work to improve our understanding of existing museum speci-
mens and to acquire the new and properly documented collec-
tions which we owe to our successors. But the new Museums
of Man must be research organizations, with the collections of
artifacts and other documents under the care of Curators of
Collections, supervised by the scientists who are supported to
do good anthropology whether or not this is directly related to
the collections. In such an enviroment we can be quite sure
that the collections will survive, that research on them will in-
crease, and that museums can significantly advance anthropol-
ogy as a whole.

Anthropology does indeed need museums. But it needs the

Natural history collection symposium 647

Very Model of a Modern Anthropology Museum, not an equiv-
ocal and petrified institution which reminds one of a bordello.”

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9TI am indebted to many colleagues for comments and criticisms which helped me
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will forgive me for not listing

they influenced me—sometimes surely insufficiently
their names.

648 Proceedings of the Biological Society of Washington

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schaft und Kunst in Deutschland. G. A. Kummer, Zerbst.

Lerris, MicHen. 1939. L’Age homme. Gallimard, Paris.

Mason, J. ALDEN. 1960. Observations on the function of the museum in
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of Canada Bull. 204, Contribs. to Anthrop. 1963-4, Pt. I,
Paper No. 4, pp. 119-127.

MercIER, PAuL. 1966. Histoire de lanthropologie. Coll. “Le Sociologue,”
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Murray, Davip. 1904. Museums: their history and their use, with a
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Palais du Trocadéro, Paris. 32 pp.

Nowotny, Kart A. 1960. Mexikanische Kostbarkeiten aus Kunstkam-
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1967b. A model for inventorying ethnological collections.
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1967c. A pilot study for inventorying ethnological collections.
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STURTEVANT, WiLLAM C,. 1966. Ethnological collections and curatorial
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TROUFANOFF, I. P. 1966. Museum of Anthropology and Ethnography of
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1967. “Grandmotherology and museology.” Curator, 10 (1):
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650 Proceedings of the Biological Society of Washington

THE ROLE OF MUSEUM COLLECTIONS IN ORNITHO-
LOGICAL RESEARCH

By Ricuarp L. Zust
Smithsonian Institution, Washington, D. C.

Periodically the curator of any large collection of birds must
ponder the future course of development of the collections in
his care in keeping with changing research needs. His plans will
probably be influenced by limitations of budget, space, and as-
sistance; and to varying degrees the development of the collec-
tion may be out of his control. The United States National Mu-
seum, for example, is a repository for birds collected for federal
agencies and as such houses rich collections of North American
birds from nineteenth century railroad surveys, boundary sur-
veys, the Biological Survey, and other sources. Today it receives
specimens from medical research units of several governmental
departments engaged in the study of arthropod-borne viruses.
Like other museums, the U. S. National Museum has acquired
large collections from privately financed expeditions. Particu-
larly in the past, the research of leading ornithologists was
molded in part fortuitously by the advent of such collections
simply because ornithological research consisted mainly of
naming new forms, reporting on new collections, writing faunal
works, and revising and classifying taxa. Even at the tum of
the century, Robert Ridgway, an esteemed curator of
birds at the U. S. National Museum, differentiated between
scientific ornithology as practiced by the curator, and popular
ornithology—the study of habits, songs, nesting and other as-
pects of life-histories. Charged with the responsibility of pub-
lishing the ornithological results of work by the government,
he produced a taxonomic synthesis of North and Middle Ameri-
can birds, based largely upon the collections of others (Ridg-

53—Proc. Bion. Soc. Wasu., Vou. 82, 1969 (651)

652 Proceedings of the Biological Society of Washington

way, 1901). This life work secured his reputation as a leading
ornithologist.

It was during Ridgway’s time, however, that study of the
living bird began to take its place with taxonomy as truly scien-
tific ornithology. Today a tabulation of research papers in a
major American ornithological journal, The Auk, would show
only about 25 percent that deal with classification and distri-
bution; an even smaller percentage of these papers are au-
thored by curators of museum collections. Although there has
been a decline in the proportion of research projects based on
specimens to those not requiring collections, the actual number
of workers who rely on specimens for their research has not
declined. Furthermore, specimens are now used not only for
traditional research activities but also in connection with other
aspects of the biology of birds. In general the research career
of a museum curator is less influenced by incoming collections
now than in the past; rather, collections are increasingly in-
fluenced by a curator’s research. Ornithology has reached the
stage where the curator, in planning the growth of his collec-
tions, must first decide what is worthwhile research.

Research on birds in museums has changed because of the
breadth of achievement of scientific ornithology as defined by
Ridgway. About 8700 species, and between three and four
times that many subspecies of birds are presently recognized;
in the past ten years only about 5 new species and some 30
subspecies have been described per year. Some of these new
species were recognized through restudy of museum speci-
mens; others were taken in the field by collectors working in
new areas—today notably within Peru and the Philippines. In
general, however, the description of new species is no longer an
important ornithological activity in that classifications are
rarely upset by the addition of the new forms. By contrast, the
description of fossil forms continues to provide new insights
into adaptive radiations of the past.

The general outlines of geographical distribution of birds are
known; indeed the final volumes of a distributional check-list
of the species and subspecies of birds of the world begun in
1931 by James L. Peters are now nearing completion (Peters,
1931). In addition to systematic and faunal treatises for most

Natural history collection symposium 653

regions of the world we now have well illustrated pocket field
guides for such areas as Argentina, Mexico, the West Indies,
North America, Europe, Russia, East Africa, South Vietnam,
Thailand, Japan, New Zealand, and the oceans. Most major
collections in larger American museums have been documented
in reports of one kind or another, and many of the larger fami-
lies of birds have received taxonomic revision in the last 50
years, some indeed receiving multiple treatment.

What then, are the current directions of ornithological re-
search? Although description of new forms has tapered off
there is still considerable interest in distributional and faunal
problems, particularly concerning South America, Africa, and
tropical Asia. In addition, systematic work continues apace with
ever-changing concepts and methods (see Sibley, 1955; Mayr,
1959). The museum worker of today thinks very differently
from his counterpart of 100 years ago and he uses such varied
approaches to systematics as comparative biochemistry, be-
havior, song structure, and functional anatomy, as well as the
more traditional ones. Museum workers and others using col-
lections have also dealt with the analysis of population varia-
tion and modes of speciation using large samples and quantita-
tive methods, with the adaptive significance of anatomical
structures, with modes of evolution on islands, with the origin
and history of avifaunas, and with a variety of other problems.
Although the limits of higher categories of birds are well estab-
lished the phylogenetic relationships of the families and orders
are little understood; these are being studied from the view-
points of comparative anatomy, behavior, and biochemistry.
There have been recent attempts to understand the adaptive
significance of the diverse patterns of reproductive biology,
migration, and molt, that have been described from the study
of specimens and living birds. Variation in characters such as
bill and wing length, measured on study skins, is playing a role
in the development of ecological theory, particularly in regard
to concepts of niche, competition, and species diversity. It is
because of the increasing emphasis on comparative biology of
birds on the part of workers who utilize specimens that we
should reexamine the nature of our specimens and collections

654 Proceedings of the Biological Society of Washington

to see that they continue to serve traditional functions and at
the same time meet new needs.

The nature of the specimen itself in part determines its use-
fulness or potential as a research tool. A typical bird specimen
is made by removing and discarding nine of the bird’s ten organ
systems, filling the remaining integumentary system with cot-
ton, sewing it up in such a way that the inner portion of the
wings can never be studied, and affixing a label. In another
kind of preparation, nine organ systems are again discarded,
leaving only the skeletal system. Skeletons are not emphasized
in most bird collections although their importance is gaining in
many museums. Still less in favor is the spirit specimen—a bird
with all ten organ systems intact and preserved in alcohol. The
early skeleton and spirit collections, often abhorrent to the skin
taxonomist, owe their existence less to ornithologists than to
workers in museum divisions of comparative anatomy. Other
specimens once of great moment but now rarely consulted are
the empty egg shells and the empty nests. All of these prepara-
tions are the traditional tools of the ornithological curator’s
trade.

For a single organ system, the bird’s skin and feathers con-
tain much information; it is this part of the bird that meets the
environment and this part to which other birds react. Feathers
are therefore subjected to many selection pressures and they
have evolved an enormous diversity of structure, pigment, pat-
tern, and molt sequences. (Because feathers of the folded
wings of a bird skin are difficult or impossible to study, the
molt pattern of the wings should be routinely recorded on
specimen labels, and some spread-wing specimens should be
prepared.) Taxonomic information from the integumentary
system applies chiefly to relationships at the infraspecific and
specific levels because differences often reflect geographic
isolation, or the need for reproductive isolation between closely
related sympatric forms. Plumage patterns are fairly stable in
some groups, thereby serving also as indicators of generic rela-
tionships.

In the bird’s skeleton the long history of common descent
within an order or family is often reflected by peculiarities of
the relationships of bones. In addition, by its proportions the

Natural history collection symposium 655

skeleton strongly reflects behavior patterns of feeding and lo-
comotion that may characterize related groups or unrelated
ecological counterparts. Comparative and functional osteology
therefore have served as useful bases for establishing the higher
taxonomic levels and for understanding structural adaptation.

Spirit specimens, like skeletons, have provided important
foundations for delineating higher taxa through comparative
anatomy (for example see Firbringer, 1888), and will probably
retain their importance for future studies of phylogenetic rela-
tionships.

In the past, skins were studied by taxonomists—skeletons and
spirit specimens by anatomists. Anatomists (and paleontolo-
gists ) had to be content with the few anatomical specimens that
were prepared along with the multitudes of skins obtained on
expeditions. Anatomists have therefore become used to work-
ing with one or a few specimens, often with incomplete data,
but many skeletons (preferably at least ten of each sex) are
needed to encompass natural variation and to avoid errone-
ous conclusions based on artifacts of preparation. Spirit speci-
mens are also needed in large series because several organ sys-
tems may be destroyed during dissection of any one system.
Some collectors “pickle” specimens that are too damaged to
skin, when in fact there is nothing more useless than a badly
damaged anatomical specimen. The data vital for skin labels
are equally vital for anatomical specimens. Today anatomists
are a vanishing breed, but many ornithologists undertake an-
atomical studies for the solution of ornithological problems. It
is increasingly apparent that many questions in avian biology
and taxonomy cannot be answered by using skin collections
alone. The traditional skin collection should therefore give way
to balanced collections for each species, including skins, skele-
tons, and spirit specimens, as well as neonatal young, eggs, and
nests. Any curator who fails to develop all of these kinds of
collections is simply limiting the research potential of the mu-
seum at a time when the need for diversity of approach to prob-
lems is rapidly increasing.

To a limited degree wholly new kinds of collections are be-
coming a part of the ornithologist’s bag of tools. The Library
of Natural Sounds at Cornell University, containing about 300

656 Proceedings of the Biological Society of Washington

miles of tape, can be called upon for comparative study of bird
songs. Files of X-rays may be regarded as supplementary col-
lections, and slide collections of comparative histology will
probably be available some day. Samples of egg whites and
blood may be stored temporarily until permanent records of
their chemical properties are made and filed. Comparative
study of birds in the field is like an extension of the specimen,
especially when documented by photographs or motion pic-
tures. Methods of storing and making available such new “spec-
imens” are in general not well developed.

Having examined the specimens let us now look at the merits
and demerits of different kinds of collections in the light of
present research trends. Collections made today or in the fu-
ture are likely to be of three different sorts: Those of a general
nature made with no biological problem in mind but intended
to increase representation of certain portions of a museum’s
holdings; collections designed for the solution of a particular
ornithological problem, collections designed for the solution of
a non-ornithological problem. The need for general collections
from all parts of the world has diminished with the advancing
development of traditional ornithology, but the need for im-
proving world-wide representation in the larger museums and
regional representation in smaller museums continues because
of the value of collections as seed sources for ideas. In other
words, although it is often possible to assemble enough speci-
mens from many museums to answer a given question, the
question might not have been asked without sufficient repre-
sentation of species or specimens in any one museum to show
that a problem existed. Important research museums should
therefore inventory their holdings and attempt to fill in gaps
within the overall scope of their collections. This job could be
done by a collection manager and trained collectors, leaving the
research curator free to specialize.

Collections intended to solve a problem, whether or not an
ornithological one, are sometimes analogous to a laboratory ex-
periment in which most of the variables are controlled. Birds
vary by age, sex, season, color phase, geographical origin, eco-
logical situation, and physiological cycle. To study the causes
and properties of any one variable the other variables can be

Natural history collection symposium 657

minimized by selective collecting. Only in a specialized col-
lection are adequate series of the critical specimens likely to
be obtained. Examples of such collections are: sibling species
taken just after their complete molt for comparison of subtle
color differences in fresh feathers; series taken across zones of
allopatric hybridization for study of gene flow in populations;
specimens taken at regular time intervals throughout the year
to determine the molt and breeding regimen of a population;
comprehensive collections from a given locality and season for
ecological or faunal comparisons with other such collections.

Examples of non-ornithological problems requiring collec-
tions are: determination of the role of a given species or re-
gional population of birds in carrying viruses or their arthropod
vectors; evaluation of the involvement of a species or local avi-
fauna in the destruction of an agricultural crop. Here the virus,
parasite, or stomach contents are the primary collections
whereas the bird specimen may be retained only for species
verification. The ornithological value of such studies could be
slight or great depending on the degree to which factors of
ornithological importance were added to the initial research
objectives.

Specimens derived from a specialized project will, in some
‘ases, be obtained in much larger series than necessary to fill in
the desired representation in the museum’s general collection.
This is particularly vexing when large birds are involved. If
space is a problem one could argue for discarding such speci-
mens at the conclusion of the study on the grounds that their
intended purpose had been served. For reasons mentioned
later I believe they should be retained or distributed to other
museums.

What sorts of data should be associated with specimens in
future collections to enhance their usefulness for research? This
question has been dealt with in different ways by others (for
example, Miller, 1940; Van Tyne, 1952; Parkes, 1963). Tradi-
tionally, the principal data recorded with each specimen has
been the locality, date, sex, and collector. Of these, the first
three, and to a lesser extent the last, are objective data that
everyone can (usually ) interpret without ambiguity. Other types
of data are often added to the label today, as they were indeed

658 Proceedings of the Biological Society of Washington

by some of the earlier collectors—skin colors, weight, amount of
fat, stomach contents, presence of brood patch, breeding con-
dition, molt, etc. Some of these data are subjective in that they
can be misinterpreted by a research worker unless they were
carefully qualified or described by the preparator. For ex-
ample, body weight may be recorded to a tenth of a gram—
but how fat was the bird, and how much did the fish in its
stomach weigh? Body feathers may be said to be molting, but
was the bird really molting or was it just replacing some
feathers lost accidentally? What unrecorded soft or liquid
foods were eaten, leaving no trace in the stomach? Does “testes
enlarged” necessarily indicate breeding and does “skull un-
ossified” indicate immaturity? Such data are of greatest use
when qualified so as to minimize their subjectivity. Subjective
data should not be confused with items such as “sex,” the deter-
mination of which requires recognition of sometimes tiny and
confusing internal organs and is therefore subject to error, but
not to interpretation.

The integrity of the specimen label determines the scientific
usefulness of the specimen and of the collection. Data on
labels are subject to errors stemming from carelessness, igno-
rance, and fraud. To reduce errors of carelessness the label
should be made out at the time of collection and preparation,
and attached to the specimen by the preparator rather than tran-
scribed from a field book by someone else later, and associated
with the specimen on the basis of a field number. Errors of
ignorance can be reduced (and subjective data enhanced ) by
training collectors in those aspects of avian biology that are
pertinent to the production of a useful label. Knowledge of
the source of data on a label can be useful to the scientist in
judging the likelihood of errors of all kinds and it follows that
the name of each person who records data must appear clearly
on the label. In collections made for non-ornithological pur-
poses or in large ornithological expeditions it is sometimes the
case that only the name of the project, or the sponsor, or the
principal investigator, appears on the label. One is then at a
loss to know who recorded the data.

In an attempt to facilitate research and curation of certain
collections, the Smithsonian Institution is developing an elec-

Natural history collection symposium 659

tronic data processing system (EDP) capable of storing, sorting
and printing out much of the data associated with specimens
(Galler, et al., 1968). The advent of computer technology
may seem to be an argument for amassing more general
collections with more data on the labels because the computer
is capable of sorting and combining voluminous amounts and
diverse kinds of information. The research value of an EDP
printout, however, is limited by the accuracy of the data and
by the difficulties inherent in subjective data, compounded by
the nagging possibility of operational errors. There is the dan-
ger that printouts of specimen data, if readily available, would
generate a rash of research based on printouts without ref-
erence to the specimens—hence without critical evaluation of
the accuracy, reliability, or significance of the data. Many
questions will not be answerable by the data selected for in-
clusion in EDP; any attempt to rectify this difficulty by record-
ing “complete” data in the field is self-limiting in that it would
leave little time to obtain and prepare specimens. Specialized
data cannot be gathered by untrained assistants. The alleged
research and curatorial values of computerizing all museum
collections are limited by these and other difficulties, and they
must be weighed against the costs (in time and money) of
setting up and operating the system. Although the practicality
of a shot-gun application of EDP to all ornithological collec-
tions is doubtful, data processing could be a useful tool in some
research projects if the data were gathered in such a way that
important questions could be answered within the capabilities
and limitations of the machine.

As a preliminary step in planning research on museum speci-
mens it would be useful to know in which museums or collec-
tions the desired specimens could be found. This need could
be most simply satisfied if each museum were to publish an in-
ventory of its holdings by species (or subspecies if possible ).
More useful would be a composite inventory of all museum col-
lections following the form of the Union List of Serials, in
which the serials would be replaced by bird species (or sub-
species ), and the libraries by museums (with a rough indica-
tion of numbers of specimens). Even an incomplete compen-
dium would be immediately useful and would gain in

660 Proceedings of the Biological Society of Washington

importance as additional museums incorporated their inven-
tories. This would be a modest undertaking compared with the
Union List of Serials, which records the representation of over
150,000 titles in 956 cooperating libraries.

In ornithology I anticipate that general collecting will con-
tinue to decline as all portions of the world become better rep-
resented by specimens, and that incoming collections will be
geared more for answering particular biological questions than
for producing conventional collection reports. I believe, how-
ever, that a balance between general and special collections
should be maintained because of the value of general collec-
tions for bringing to light the unexpected.

Curators will have to decide whether or not to retain special
bird collections that have served their purpose in answering a
particular problem. In deciding we must remember that col-
lections cannot be duplicated with the ease of a chemist dupli-
cating a precipitate and that their research potential surely ex-
ceeds that realized in any one study. In some respects their
usefulness increases with time; over a period of 50, 500, or
1000 years specimens may, like fossils, provide the evidence for
evolutionary change and rates. (After all, what are fossils but
skeleton collections that have been housed in rock rather than
boxes?) Also, specimens become important historical docu-
ments as particular environments on earth are changed or
lost. Another reason for retaining collections is that the pub-
lished word represents opinion and is subject to error; as ideas
change and as the literature becomes distrusted after a period
of years reference to the specimen is required again (Berlioz,
1960 ). If a museum cannot provide accessible storage space for
increasing collections an effort should be made to distribute at
least parts of long series to other museums or to teaching in-
stitutions.

Emphasis in museum ornithology will probably remain for
some time on various aspects of the comparative biology of
birds and on the processes of speciation and differentiation of
the higher categories
one working in behavior, ecology, ecological physiology, cyto-

problems that may be served by some-

genetics, or biochemistry as well as in more traditional aspects
of systematics. To justify occupying a museum position, how-

Natural history collection symposium 661

ever, a curator should apply his interests toward understanding
the diversity of birds through comparative studies that in some
way derive support from collections. Diversity of approach to
collections may be the key to continued viability of museum
research as we expand from traditional functions into com-
parative biology.

LITERATURE CITED

Berwioz, J. 1960. Le role capital des musées dans l’avenir de lorni-
thologie. Proc. XII Internat. Ornith. Cong., 1958, Vol. 1, pp.
44-49,

FURBRINGER, MAx. 1888. Untersuchungen zur Morphologie und Systema-
tik der Vogel. Vols. 1 and 2. T. J. van Holkema, Amsterdam.

GALLER, SIDNEY R., ET AL. 1968. Museums Today. Science, Vol. 161, No.
3841: 548-551.

Mayr, Ernst. 1959. Trends in Avian Systematics. Ibis, 101( 3-4): 293-
302.

Mititer, ALDEN H. 1940. Field Techniques in Collecting for a Research
Museum. Museum News, 17( 17): 6-8.

PARKES, KENNETH C. 1963. The Contribution of Museum Collections to
Knowledge of the Living Bird. The Living Bird, Second An-
nual. pp. 121-130.

Peters, JAMES L. 1931. Check-list of Birds of the World. Vol. 1. Cam-
bridge, Harvard University Press (12 of 15 projected volumes
of this series have been published ).

Ripcway. Rosertr. 1901. Birds of North and Middle America. Bull. U.
S. Nat. Mus., No. 50, Pt. 1 (8 parts were published by Ridg-
way and 3 more after Ridgway’s death).

SIBLEY, CHARLES G. 1955. Ornithology. In A Century of Progress in the
Natural Sciences, 1853-1953. California Academy of Sciences,
San Francisco.

Union List oF SERIALS IN LIBRARIES OF THE UNITED STATES AND CANADA.
Third Edition. New York, The H. W. Wilson Co. 1965.

VAN TYNE, JOSSELYN. 1952. Principles and Practices in Collecting and
Taxonomic Work. Auk, 69(1): 27-33.

662 Proceedings of the Biological Society of Washington

MALACOLOGICAL COLLECTIONS—DEVELOPMENT
AND MANAGEMENT

By JoserpH ROSEWATER
Smithsonian Institution, Washington, D.C.

As a curator in the Recent mollusk section of a large museum,
one is often confronted with the question from interested visit-
ing members of the public: “How many shells do you have
here in all these cabinets?” One usually reacts by recalling the
figure cited for Mollusks in the most recent annual report:
“10,058,888” or “somewhat in excess of 10,000,000.” The prop-
erly impressed visitor often counters with the predictable
phrase: “Oh my, but where do they all come from?”

Before going into the answer to that question, at the outset,
I should like to emphasize a fact which I believe is largely lost
to many persons who like to think “collectively” about collec-
tions. Collections of shelled mollusks are different from many
other kinds of natural history collections. A living shelled mol-
lusk is just that; it comes in at least two major parts, the shell
and the soft animal which forms the shell. And mollusk col-
lections are almost universally curated in at least two parts also:
(1) a collection of dried shells, arranged usually according to
the most accepted classification and (2) a collection of pre-
served soft animals, maintained separately from the dry shells
and arranged so as to provide some means of rapid access for
study in conjunction with the shells.

This system of collection storage in which two collections
actually are maintained may appear unwieldy to many, and it
is true that it has come down to us from the last century, al-
though altered in many ways as new materials and methods of
handling and storing mollusks came to light. Because the sys-
tem is old, however, does not mean by definition that it is bad,
nor impossible to work with, nor that it should be changed im-

54—Proc. Bron. Soc. Wasu., Vou. 82, 1969 (663 )

664 Proceedings of the Biological Society of Washington

mediately. It means that it is the most satisfactory compromise
reached so far for the maintenance of a collection of important
animals. By common agreement of Malacologists, the mollus-
can exoskeleton or shell is the most important single diagnostic
tool for the discrimination of species. The collateral softparts
contribute additional information concerning species identity
and supraspecific relationships.

The collection of dry shells is somewhat unique among cate-
gories of biological specimens, although obviously it has much
in common with certain other groups of invertebrates, and
strangely enough, mammals and birds and, of course, minerals
and fossils. But the shell is often a very durable object which
endures handling and storage much better than the majority of
other specimens. Its catalogue number may be written directly
upon it, as well as upon associated labels containing habitat
and locality data. Modern shell collections are usually stored
directly according to their classification, beginning at one end
with the most “primitive” forms and proceeding through the
more “advanced” ones. I have often heard the question: “Do
you have a card file of the species present in your collection and
perhaps a cross-reference file to their geographic data?” No,
we do not

and it would be nice to have such a reference, es-
pecially one provided by a computer by which specimen data
could be extracted in all imaginable ways and correlated. Per-
haps when classification of mollusks reaches a_ sufficiently
stable point, it will be profitable to enter their data into such
a system. At present, only certain groups are ready for ADP.
But we do not make lists of the collection, because the collec-
tion is arranged in such a way as to form its own list and spe-
cies may be arranged geographically within this system. One
who is familiar with the classification of mollusks can work
easily with such a collection after a brief orientation based on
individual collection idiosyncrasies.

Mollusks differ also from many other groups in the degree of
interest with which they are favored by the layman or amateur.
“Breathes there the man (or woman, or child) with soul so
dead” that he has never stooped down and picked up a shell
during a visit to the seashore? It may be added that a goodly
percentage of these persons carry their shells straight to the

Natural history collection symposium 665

curator of mollusks for identification! It is doubtful that any
other group of animals is so widely collected by man. Their
popularity has often brought on invidious comparison to stamp
collecting, another extremely well participated hobby through-
out the years.

These introductory remarks have been intended to place in
proper prospective the mollusk collection. To the professional
malacologist, it is a vehicle for his research, often oriented
toward systematic and zoogeographical pursuits. Few groups of
mollusks are adequately understood as yet. Some, especially
commercially productive ones, have been thoroughly explored
and their biology and classification are well known. But tens-of-
thousands remain to be studied, and the most feasible way to
study the systematics and distribution of these animals is
through large series in museum collections.

The question was posed earlier: where did our collections of
mollusks come from, that is, how did they develop. In discuss-
ing the development of Malacological Collections, I shall limit
myself to only a brief consideration of collections outside of
the United States, but since collections began abroad, we must
start there.

In the writings of Aristotle we find considerable mention of
mollusks and so it is apparent that they were important objects
of man’s interest at a relatively early date. Shells unearthed
from the rubble of Pompeii indicate a collection of some sort
had been put together there, not only of specimens from the
Mediterranean, but from the Indo-Pacific region. Cicero’s writ-
ings make mention of shell collecting as a relaxation from the
tribulations of war and government. It is said that the first
large-scale expedition in search of shells took place in 40 A.D.
when Caligula led his troops down to the sea in Gaul as if to
embark on an invasion of Britain; having drawn them up in
battle formation, he ordered them to collect shells—which he
called ‘the spoils of conquered ocean.’

Our knowledge of the development of Malacology during the
Middle Ages, as it is with so many branches of knowledge, is
limited to the literature produced in the monasteries. Some
quite recognizable species were illustrated in the exquisite il-
luminated manuscripts of that day.

666 Proceedings of the Biological Society of Washington

With the coming of the Renaissance and the age of dis-
covery, natural history cabinets flourished throughout Europe,
usually in the hands of rich men who had the time and finances
to accumulate such collateral wealth. During this period the
first small museums of natural history specimens came into
being and some of their most popular contents were well known
to have been the shells of mollusks. These collections often
were accumulated by or found their way into the hands of
noblemen who enlarged them and saw to it that they were con-
served. By the 17th and early 18th centuries several of the royal
houses of Europe had amassed large collections of shells. The
celebrated Linnaeus was commissioned by the queen of Swe-
den to arrange her shells and upon her collection are based
a number of the molluscan species in the 10th edition of “Sys-
tema Naturae.” Thus, gradually through the assimilation of
small and private shell collections by the rich and by royalty,
rather massive holdings were acquired which eventually had
established for their conservation the Natural History Museums
which we know today.

Concerning the development of Malacology in the United
States, I should like to quote from an address made by William
Healey Dall to the Biological Society of Washington at its 8th
Annual Meeting 80 years ago in 1888. Dall said, “I may divide
the study of Mollusca in this country into three periods, al-
though these are connected by many intermediate links. The
infancy of the science, with a Linnaean classification, has no
representation in American conchological literature, which
sprang, full-grown, like Minerva from the head of Jove, from
the Lamarckian school of Europe. The first period might fitly
bear the name of its inaugurator, Thomas Say. It is character-
ized by a rapid advance in the determination of the fauna, the
classification of the species, and the exploration of vast areas.
It extended from 1817 to 1841.

“The second period should bear the name of Dr. A. A. Gould.
It was inaugurated by his report on the Invertebrata of Mas-
sachusetts (1841), and characterized by the broader scope of
investigation, and interest in geographical distribution, the
anatomy of the soft parts, and the more precise definition and

Natural history collection symposium 667

exact discrimination of specific forms, as exemplified in his
writings.

“The third period would be appropriately called after Dr.
William Stimpson, who eagerly adopted the radical changes in
classification rendered necessary by the discoveries of Loven,
and [who] stood ready to welcome the theory of evolution with
all the light it shed in dark places.”

The name of Thomas Say is much revered in American Mala-
cological circles. He is called the “father” of that branch of
science in this country, and was early associated with the first
institution in our country to boast a collection of mollusks, The
Academy of Natural Sciences of Philadelphia, established in
1812. There were many natural history societies in the years
that followed, small local groups of persons who gathered to
discuss, collect and study various facets of our new country’s
natural history. In New England, the Boston Society of Natural
History superseded the Linnean Society in the early 1830's. The
Smithsonian Institution made its appearance in the middle 19th
century with an “instant” mollusk collection accumulated as a
result of the U. S. Exploring Expedition. Data gathered for a
history of the Division of Mollusks of the U. S. National Mu-
seum by Dr. Harald A. Rehder show the Smithsonian collection
of mollusks had its beginning as early as 1840 with the organiza-
tion of the “National Institution for the Promotion of Science”
established in part as a repository for the Exploring Expedition
collections. In 1860 the Agassiz museum in Cambridge opened
with the beginnings of a mollusk collection which would one day
absorb the specimens brought together by the Boston Society
of Natural History as well as many large private collections.

Of the several large museum mollusk collections of this
country today, four of the largest are located in the east: they
are at the Museum of Comparative Zoology, the American Mu-
seum of Natural History, the Philadelphia Academy and the
Smithsonian Institution. Others are at the Museum of Zoology,
University of Michigan, the California Academy of Sciences,
San Francisco, at Stanford University and at the Los Angeles
County Museum. The large collections maintained in these mu-
seums are partly the result of specimens returned by expedi-
tions mounted wholly or in part for the purpose of collecting

668 Proceedings of the Biological Society of Washington

natural history specimens. But they are in large part also the
result of the donations by individuals of anywhere from single
specimens to entire private collections consisting of thousands of
specimens. And so it was that my original example, the Smith-
sonian collection of mollusks, came to be estimated to number
in excess of 10 million specimens.

I will base my remarks on management of Malacological
collections upon my general knowledge of these practices
gained through association with the Smithsonian-Division of
Mollusks. The task of managing or curating the largest col-
lection of mollusks in the United States, if not in the world,
has not been something to be faced by any one curator, for ob-
viously the collection has developed through time. At one
time the standard procedure for preparing the shells was to
glue them onto glass plates or cardboard or wooden plaques
and to inscribe these with the names and other data. Rehder’s
manuscript history describes how Dall, who as the first virtual
curator of mollusks, inherited the collection so prepared for
the Smithsonian by P. P. Carpenter. He struggled with these
mounted specimens, remounting them as they fell off their
plates. Dall, prodded by this unwieldy and space consuming
curatorial procedure, finally removed the specimens from the
plates and placed them in vials and small trays, each lot with
its data-containing label, thus instituting the space conserving
procedures used today.

With today’s vastly improved transportation more and more
persons are getting into the field—more and more both large
and small expeditions are being mounted and many of these
are bringing back mollusks. Over the past twenty years we
have managed to accession an average of 58,000 specimens per
year, over a million altogether, the real total of specimens re-
ceived being in excess of that figure because some collections
have not yet been accessioned. On this basis we may plan to
expand our collection by approximately 12 percent every 20
years if we keep constant the rate at which material is coming
in. Given the personnel and equipment for processing, cata-
loguing and storage this is not an overwhelming addition to
keep up with. But it must be stressed that the rate of addition
seems to be on the increase.

Natural history collection symposium 669

We hear a great deal these days that space available for
collections is finite and that steps must be taken to fit material
into present space. What can be done in the case of mollusks
to help the space situation and still maintain an optimum of
systematic and geographic coverage and an ample biological
series for comparison of morphological variation?

1. Curatorially for many years our dry collection has been
at the forefront of any of its size that I have seen. As new
material is added to the collection the classification is con-
stantly being updated and old material, which lacks data or
which is in poor condition and was kept only because it was
the sole example of a species is weeded out. In this way a
surprising amount of “bulk” is removed from the collection,
making room for pertinent material. Also, as groups of mol-
lusks are critically reviewed during monographic work, their
curation is brought up to date. Collections of dry mollusks are
admirably adapted for concise storage and anyone examining
the Smithsonian collection will find that it contains an enor-
mous amount of material very compactly stored.

2. In addition to careful curatorial procedures some selectiv-
ity must be practiced in the acceptance of material for the
collection initially and in the retention of specimens already
received. At one time we felt that we were compelled to ac-
cept and retain almost any collection thrust upon us. The
sheer weight of collections which have been known to ac-
cumulate in what might be called “such indiscriminate ac-
cepting’ has shown this to be a mistake. We like to be asked
but retain the right to say “no”! Then too, during the process-
ing of large collections, it is often expedient to reserve a por-
tion of many lots for profitable exchanges with other institu-
tions, in this way reducing the bulk to our own collection.

3. A third way of controlling to some degree the sorts of
material received for the collection is through specialization.
This may take the form of limiting oneself to a particular class,
order, or smaller group or by limiting the geographic area of
one’s major interest. The tendency in Malacology today is to
specialize, although those of us who received their training in
the “Old School” are used to working in two or more very dif-
ferent major groups more or less interchangeably, for instance:

670 Proceedings of the Biological Society of Washington

gastropods and bivalves. A division of labor in the phylum,
with responsibilities spread among several curators can make
feasible a more efficient curatorial team so long as their goals
are somewhat similar. Geographic specialization is practiced to
some degree in collections of mollusks. For instance, the Smith-
sonian collection has lately emphasized the Indo-Pacific faunal
area, whereas the MCZ favors the western Atlantic. Both in-
stitutions, nevertheless, try to maintain a collection which is
balanced and can be utilized for world-wide studies.

There are doubtless many other ways to exert a conscious in-
fluence over the development of a museum collection of mol-
lusks, but the preceding three: Careful curation, selectivity and
systematic or geographic specialization seem to me to be the
most natural ones and they avoid the process of subjectively
eliminating large portions of material to make way for others.

For the immediate future I can see a need for the continuance
of an orderly accumulation of additional material to collections
of mollusks. At the same time, I feel strongly that we should
have farthest from our minds the concept that mere accretion
is an end in itself. We need to study this material and create a
classification which in time may sort itself out to being some-
thing near a “natural order.” Until this is done I am of the
opinion that we will do well to bear in mind two quotations
credited to G. Brown Goode in 1895. I think they balance each
other nicely. The first: “Curators are apt to err on the side of
saving too much”; the second: “A finished museum is a dead

museum.”
REFERENCES

Dati, WitttAM H. 1888. Some American Conchologists. Proceedings
of the Biological Society of Washington, vol. 4, pp. 95-134,
portraits.

Dancer, S. Peter. 1966. Shell Collecting, An Illustrated History. Faber
and Faber, London. 344 pp., 35 pls., 31 figs.

Goutp, Aucustus A. 1841. Report on the Invertebrates of Massachu-
setts. Cambridge. xiii + 373 pp., 213 figs.

KetLocc, Remincron. 1946. A Century of Progress in Smithsonian
Biology. Science, vol. 104, pp. 132-141.

Tryon, Georce W., Jr. 1862. A Sketch of the History of Conchology in
the United States. American Journal of Science and Arts, vol.
33, pp. 13-32.

AUTOMATION IN MUSEUM COLLECTIONS!

By RaymMonp B. MANNING
Smithsonian Institution, Washington, D. C.

INTRODUCTION

For a little over a year now several of us in the Smithsonian
have been associated with a project designed to investigate
possible uses of electronic data processing (EDP) for computer
storage and retrieval of specimen-associated data. The project
has been funded under a contract with the Office of Education
of the Department of Health, Education, and Welfare. It is a
joint effort by members of the staff of the Museum of Natural
History and Information Systems Center, Smithsonian Institu-
tion.

The project is based on the thesis that a museum collection
is more than an assemblage of inanimate objects or dead orga-
nisms; it is a vast information resource which we cannot ade-
quately use with current methods of record keeping. A second
factor, which is also quite important, is that collections are con-
tinually growing at a rapid rate. In the Department of Inverte-
brate Zoology alone, the collections are increasing by at least
200,000 specimens per year. This trend is hardly likely to change
in the near future, and if specimen-associated data in the
collections is too difficult to obtain now, it will be even less
available in the future. If computerized data record-keeping
systems are going to be developed and used, the project must
be started now. Delays will only increase the difficulties and
the cost.

The MNH project was designed to set up record-keeping sys-
tems in three separate areas of the museum: marine rocks, un-

1 This work was supported by a grant from the Library and Information Sciences
Research Branch, Office of Education, Department of Health, Education, and Wel-
fare, OEG-1-071159-4425.

55—Proc. Biot. Soc. WasH., Vou. 82, 1969 (671)

672 Proceedings of the Biological Society of Washington

der the supervision of William Melson, Department of Mineral
Sciences; oceanic birds, under the supervision of George Wat-
son, Department of Vertebrate Zoology; and marine crus-
taceans, particularly stomatopods, under the author's supervi-
sion. Donald F. Squires, then Deputy Director of the Museum,
was among the first to recognize the need for data processing
in the museum and it was he who sought and obtained support
to initiate the pilot project.

In Crustacea, the project is designed to aid management of
the collection, to aid curation, and to enhance the collection as
a research tool. Our primary aim was to update some of our
techniques of collection management, and to develop tech-
niques which would allow us to manipulate specimen-associ-
ated data without having to return to the collection every time
we work with the data.

The overall MNH project has been divided into two phases:
(1) to build a data bank based primarily, but not exclusively,
on three separate collections, and (2) to manipulate the data
in various ways to evaluate the overall costs of not only various
portions of the project but also the costs of general handling
and processing of museum collections, regardless of the ulti-
mate use and method of storage and retrieval of the data.

The first 18 months of the contract, ending in December,
1968, have been concerned with entering the data, building up
the base, and solving the innumerable problems that arose at
every step. The next phase will deal principally with interroga-
tion of the data base.

My remarks are designed to give a progress report on activi-
ties in Crustacea, an idea of some of the different problems
which we have encountered and some of the results of the proj-
cr.

We are not alone in the scientific community in our interest
in developing a system for storage and retrieval of specimen
data. The British Museum ( Natural History) and the National
Museum of Canada are both working on developing such a
system, and system development is being considered by over
30 museums around the world. More than 70 representatives
of universities, museums, zoological parks, and botanical gar-

Natural history collection symposium 673

dens met in Mexico City, in 1967 to discuss “Information Prob-
lems in the Natural Sciences.” Bullis and Roe (1967), report-
ing on a bionumeric code used by the Bureau of Commercial
Fisheries Exploratory Fishing Base, Pascagoula, Mississippi,
noted that faunal data resulting from their exploratory fishing
operations necessitated development of computer methodology
for handling the data.

Those of us associated with the MNH project are not the only
ones in the museum working with different applications of data
processing. In the Department of Botany, Stanwyn Shetler is
applying EDP to the broad Flora of North America Project,
and Mason Hale is developing a type-catalog, and a list of gen-
eral accession records and invoice data in Botany have been
computerized for some time. James Peters, Department of
Vertebrate Zoology, has pioneered within the museum in devel-
oping computer programs to carry out time-consuming statisti-
cal analyses commonly used in taxonomic studies.

In a recent article on curation of invertebrate collections,
Emerson and Ross (1965, p. 337) noted that: “The ideal method
for cataloguing specimens and the retrieval of catalogue and
specimen information would be a punch card or magnetic tape
system. Vast amounts of information could be stored in a
relatively small space and retrieved within seconds. Unfortu-
nately, none of the museums in the United States has yet in-
stalled such a system.”

BACKGROUND

The Division of Crustacea was most fortunate to receive
through the foresight and industry of members of its fore-
runner, the Division of Marine Invertebrates, a remarkable
file of specimen records in a 3 X 5 card format. This card file,
consisting of about 125,000 entries, serves as a guide or index to
the collection, a source of information on loans and holdings in
general (management data), and a basic source of specimen-
associated data. The existence of the record file, in a museum
where specimen records, other than specimen labels and cata-
log books, are in general absent, is a tribute to the persever-
ance and foresight of such people as Mary Jane Rathbun,
Waldo L. Schmitt, and Fenner A. Chace, Jr. The Marine In-

674 Proceedings of the Biological Society of Washington

vertebrate catalog system has existed for approximately 70
years.

Our new system in Crustacea was designed as closely as
possible to that already in existence in the Division. We wanted
to show that we could continue the basic operations of docu-
mentation and cataloging of the collection and prepare the
data for computer storage at the same time.

Briefly, let me summarize the pre-computer method of cata-
loging. A cataloger would compile the data necessary for each
entry, hand enter it into a ledger catalog, hand-write the label,
type two copies of a specimen data card (one copy was to be
filed in the species file, the other to be filed in a geographic
file), and type a neck label for the jar.

In 1965, we instituted a change in the cataloging procedure
by installing a typewriter system in which a punched-paper
tape could be generated during the initial typing of the data.
The machine used is a CDC (SCM) Typetronic 2816 with two
typewriter consoles, one featuring microelite type (with 16
characters to the inch) and one featuring standard elite type
(12 characters to the inch). As the data is entered on the speci-
men label on the microtypewriter, the machine generates a
punched paper tape which can then be used to reproduce
automatically on the other typewriter as many 3 X 5 cards as
needed for the files. The jar neck label also can be typed from
the tape. The system was developed with the expectation that
some day the data on the paper tapes could be converted to
magnetic tape, but it was installed almost two years before we
received support for the MNH project.

The cards, labels, and neck labels are printed in long per-
forated strips which are much easier to feed into the typewriter.
Pink cards and distinctly-marked neck labels are used for types.
Originally, we planned on printing up three sets of 3 x 5 cards,
one for the species file, one to be filed in numerical order as a
replacement for the permanent ledger catalog, and one for the
geographic file. Henry B. Roberts, Senior Museum Specialist in
the Division of Crustacea, did most of the work involved in
developing the new card format from the old one.

At the time the program was started, Smith-Corona-Marchant
(SCM) (now Control Data Corporation) was the only manu-

Natural history collection symposium 675

TaBLeE 1. Data Organization, Division of Crustacea.

Field Maximum
Name Length
Nomenclature type 15 spaces
Catalog number 8 "
Genus name Di. "
Subgenus name, if used 21 "
Species name 21 "
Subspecies name, if used i "
Author 50 "
Total number of specimens 5 "
Location I: Continent, country, ocean 30 "
Location II: State, province, island group 30 "
Location III: County, parish, small island 30 "
Location IV: City, lake, miscellaneous 70 Ft
Latitude and Longitude 48 "
Collection Gear 20 "
Depth 20 u
Collector 20 "
Collector’s number 12 "
Date of collection 11 "
Identifier 33 "
Date of identification Gl "
Number and sex of specimens 45 "
Accession number 10 "
Type of entry (gift, etc. ) 13 "
Publication information 45 "
Preservative 3 "
General remarks, and overflow from 180a 45 "
General remarks 45 "
General remarks 45 "
General remarks 30 "
Data cataloged 11 "

facturer of a system with a micro-elite typewriter. Since then,
we have learned that SCM no longer will manufacture the
Typetronic. Fortunately, perhaps, for those who require the
micro-typewriter, Friden now manufactures one and can supply
a system comparable to the Typetronic.

A more detailed account of the development and use of the
cataloging procedure was given by Squires (1966).

676 Proceedings of the Biological Society of Washington

USNM- DEPARTMENT OF INVERTEBRATE ZOOLOGY PARATYPE

cAT- NO. 100932 ® Lysiosquilla grayi Chace ddan
@e@e0@ 2 SPECS. 7 @0e@6
LOCAL. eeee
cece United States;Massachusetts;Cape Cod; Bass ecce
coco @@800
Ce ee) DEPTH intertidal @@e0e
COLL. BY Gray, M B. 18 Mar 1957 e008
DET. BY Chace, F. A. dre TT ddd
NO/SEX 2g‘, 5

ACC. NO. 206768 ENTERED AS PRES. ale
REMARKS

Biological Bulletin, Woods Hole, vol. 114,
Nose. 2, pe 141, pl. dy Pies. 1-5, 1958.
Muddy sand at low water.

SI-MNH-172-REV. 9-20-67 DATE CAT 14 Nov 1957

USNM-INVERTEBRATE ZOOLOGY PARATYPE
100932 . Tysiosyquilla grayi Chace

S specs. 7

United States;MassachusettssCape Cod$
Bass River/

BERTH intertidal
Gray, Me Be 18. Mar 1957
Chace, FeAcgJdre

22 c22 coece

FicurE 1. Catalog card and corresponding label used in Division of
Crustacea, Smithsonian Institution.

Types oF Data ENTERED

In our cataloging operation in Crustacea, the basic unit is a
lot; each lot contains one or more specimens. Basic data for
each lot are collected and verified by a cataloger who may re-
ceive the lot with no more data than name, number and sex of
specimens, identifier and date identified, station number and
vessel, and accession number. These data are expanded by the
cataloger to include as much of the information shown in Table
1 as possible.

Each of these types of information must be entered on the
original 3 x 5 card (Figure 1) which we retain in the division
file. In Crustacea we use data assigned to 30 different fields;
often some of these items are left blank. In the experiment on
birds some 39 fields are used and in minerals approximately 140
fields have been identified.

LYS ITOSQUILLA
CRUSTACEA

O10A

GRAYI

Natural history collection symposium

ACCESSION

NOMENCLATURE TYPE

677

100932

PARATYPE

020A MUSEUM ABBREVIATION USNM

020A CATALOG NUMBER 0100932

O35A AUTHOR CHACE

040A NUMBER OF SPECIMENS 00007

OS1A MAJOR LOCALITY UNITED STATES

052A SECONDARY LOCALITY MASSACHUSETTS

053A SPECIFIC LOCALITY CAPE COD/BASS RIVER

080A DEPTH 00000 METERS, VARIANCE 0 METERS GIVEN AS INTERTIDAL

090A COLLECTOR GRAY, S.

110A DATE OF COLLECTION 18 MAR 1957

120A IDENTIFIER CHACE, F. A. JR.

130A IDENTIFICATION DATE — -- =--- ----

140A NUMBER AND SEX MALE MT Mere M JV FEMALE F OV FV JV LARVAE
2 3:

150A ACCESSION NUMBER USNM 206768

160A TYPE OF ENTRY

170A PRESERVATIVE ALC

180A PUBLICATION INFO BIOLOGICAL BULLETIN, WOODS HOLE, VOL. 114,

190A REMARKS I NO. 2,.P. 141, PL. ‘1, FIGS. 1=5, 1958

200A REMARKS II MUDDY SAND AT LOW WATER

230A DATE CATALOGUED 14 NOV 1957

Ficure 2. Work-in-progress listing for same entry as shown in Figure 1.

Our limiting factor here is perhaps the number of characters
we can enter on a3 X 5 card. The system we use has the capa-
bility of storing some 4000 characters per catalog entry; we
have used less than 800 in developing the data card in Crus-
tacea.

The first two lines include the basic information pertaining
to that log, catalog number, name, and total number of speci-
mens; these are the initial data used by those who work with
the files and are coincidentally the basic invoice data.

Data on the card down to and including the “Determined by”
level on the card also appears on the specimen label. The re-
mainder of the information appears in the ledger catalog and
on the card, but not on the label. The label for this same lot is
also shown in Figure 1. Card size was determined by the exist-
ing files and label size was determined by the size of our basic
specimen vial for smaller specimens.

Upon completion of our cataloging process for a series of
specimens, the tape generated by the Typetronic is forwarded to
the Computer Center where the data are converted by the com-
puter to magnetic tape. Then the data items are reshuffled by
the computer to produce a Work-In-Progress Listing (WIP), a
preliminary printout (Figure 2). We use this now for a second
proofing of the original entry. The machine will automatically
mark several kinds of errors, including erroneously marked
fields, fields with no data, spelling errors in the data, data in
wrong fields, etc.

678 Proceedings of the Biological Society of Washington

Corrections may be made at any of the steps in the catalog-
ing process. The punched paper tape can be corrected after
the card and label have been proofread, and corrections of
data on the WIP listings can be keypunched to update the data
in the computer.

We have the potential of retrieving data items by field or by
any combination of fields. Further, the data can be rearranged
by fields in any format which one might require.

Note that depths are converted to meters; if a depth range is
given in the original entry, the midpoint is the first depth given
in the printout. In printouts, the midpoint (in meters), the
range (in meters) and the original entry, as given in feet,
meters, or fathoms, are all reproduced. All depth conversions
are automatic.

We have left several fields vacant in the section entitled
REMARKS. These can be used for habitat information, for ref-
erences to field notes or color photos, and so on. These fields
are unrestricted at the present time.

The basic card system in Crustacea is extremely adaptable.
Although designed for marine organisms, it has been modified
with little effort to include crayfishes where locality data may
be centered on drainage system and where information on as-
sociated species, cross-referenced to field notes, is required. We
are now adapting the format to free-living marine nematodes
and to cephalopods.

I want to discuss here in a little more detail two of the types
of information entered and used, Nomenclature and Geography.

NOMENCLATURE

The system is designed so that a taxon is a focal point for
entry and retrieval of data. For each group a master taxa list
must be compiled and entered.

In preparing the master list for the stomatopods, I have in-
cluded major synonyms. The entries for one species, Odonto-
dactylus brevirostris (Miers), are shown in Figure 3.

Data on this species can be requested by using any of the
synonyms or the senior synonym in the query. Few of our col-
lections are up to date nomenclatorially for few of us have the
help required to keep up with the name changes. By being able

Natural history collection symposium 679

000 0002 330 G ODONTODACTYLUS GENUS
000 0062 331 RG BIGELOW, 1893 REMARKS
000 0002 340 B ODONTODACTYLUS BREVIROSTRIS

000 0002 350 RB (MIERS, 1884) REMARKS
000 0002 355 SB GONODACTYLUS BREVIROSTRIS SYNONYM
000 0002 356 RB MIERS, 1834 REMARKS
000 0002 360 SB GONODACTYLUS HAVANENSIS SYNONYM
000 0002 361 RB BIGELOW, 1893 REMARKS
000 0002 370 SB ODONTODACTYLUS HANSENIT SYNONYM
000 0002 371 RB POCOCK, 1893 REMARKS
000 0002 380 SB ODONTODACTYLUS LATIROSTRIS SYNONYM
000 0002 381 RB BORRADAILE, 1907 REMARKS
000 0002 390 SB ODONTODACTYLUS SOUTHWELLI SYNONYM
000 0002 391 RB KEMP, 1911 REMARKS
000 0002 400 SB ODONTODACTYLUS NIGRICAUDATUS SYNONYM
000 0002 401 RB CHACE, 1942 REMARKS
000 0002 410 B ODONTODACTYLUS CULTRIFER

000 0002 420 RB (WHITE, 1850) REMARKS
000 0002 421 SB GONODACTYLUS CULTRIFER SYNONYM
000 0002 422 RB WHITE, 1350 REMARKS
000 0002 430 B ODONTODACTYLUS HAWATTIENSIS

000 0002 440 RB MANNING, 1967 REMARKS

Ficure 3. Synonymy of Odontodactylus brevirostris (Miers) as stored
in the computer Directory of Names.

to identify and label synonyms, entries in the data bank under
any of the names can be retrieved.

A separate directory of names is maintained and, as catalog
entries are added, the names are checked against the directory;
entries accompanying names not in the directory are rejected as
are entries under misspelled names.

Neither the user nor the cataloger need be familiar with the
numericlature used by the machine; only knowledge of the
nomenclature is required.

We have also developed a hierarchical classification, for we
believed that data must be retrievable not only at the specific
level but at any of several taxonomic levels. In Crustacea we
have compiled a hierarchy down to suborder, to which we can
eventually add families, genera, subgenera, and species. A
portion of the crustacean hierarchy is shown in Figure 4. The
numbers on the right are the numericlature; those on the left
are part of a sequence of numbers required to enter the data
originally and are not related to the numericlature.

Our numerical code of 26 digits was developed to allow maxi-
mum flexibility in adding to the hierarchy at any level and to
maintain the specific name as the key to entry and retrieval of
data. Neither the cataloger nor the scientist user needs to
know the entire number sequence; it is internal in the com-
puter.

680 Proceedings of the Biological Society of Washington

SEPT 26, 1968

GROUP
PAGE -LINE

0000029125
0000030130
0000031140

0000032150
0000032160

0000033170
0000034200
0000034205
0000034206
0000035209
0000036220
0000037230
0000037235

0000039250
0000039260

0000040270
0000040280
0000040290
0000041300

0000042310
0000042320

0000042325
000004 3330
0000044 340
0000045350

0000046358

HIGHER TAXA NUMERICLATURE LISTING

TYPE

LVL

C

ie)

Cc

TAXA

LEPTOSTRACA

NEBALIACEA

RHINOCARINA

CERATIOCARINA
CERATOCARINA

NAHECARIDA

SYNCARIDA

ANAS P IDACEA

BATHYNELLACEA

PERACARIDA

THERMOS BAENACEA

SPELAEOGRIPHACEA

MYS IDACEA

CUMACEA
SYMPODA

TANA IDACEA
CHELIFERA
ANISOPODA
ISOPODA

AMPHIPODA
LAEMODIPODA

EUCARIDA

EUPHAUS IACEA

PYGOCEPHALOMORPHA

DECAPODA

HOPLOCARIDA

NUMERICLATURE

1

1

1

1

1

032

032

032

032
032

032

032

032

032

032

032

032

032

032
032

032
032
032
032

032
032

032

032

032

032

008

008

009

O14

000

O01

000

000
000

000

000

O01

002

000

OOL

002

003

004
004

005
005
005
006

007
007

000

OOL

002

003

032 015 000

0000

0000

0000

0000
0000

0000

0000

0000

0000

0000

0000

0000

0000

0000
0000

0000
0000
0000
0000

0000
0000

0000

0000

0000

0000

0000

Ficure 4. A portion of the crustacean classification used

project.

0000 000000

0000 000000

0000 000000

0000 000000
0000 000000

0000 000000

0900 000000

0000 000000

0000 000000

0000 000000

0000 000000

0000 000000

0000 000000

0000 000000
0000 000000

0000 000000
0000 000000
0000 000000
0000 000000

0000 000000
0000 000000

0000 000000

0000 000000

0000 000000

0000 000000

0000 000000

00

in the MNH

The hierarchy, to order, was arbitrarily selected and entered
into the computer. Aspects of classification can be updated at
any time and categories below order can be entered at any
time. The classification of invertebrates proposed by Black-
welder (1963) was used as the basis for the hierarchy.

Assume that we have completed sections in Crustacea and
now plan to enter any other invertebrate group. The hierarchy

Natural history collection symposium 681

will accommodate 999 families under any order, and, further,
up to 9 additional families can be entered at some future time
between any 2 of the original entries, [without changing the
numericlature assigned to the original families entered]. Simi-
larly, up to 99 genera can be assigned to each family and 9 gen-
era can be added between any of two of the original genera.
The number of species that can be entered originally in each
genus is 9999, with room to add 99 between any 2 of the orig-
inal entries.

Data can be entered by genus or higher category only and
also be retrieved by those categories. Data associated with
such designations as variety, forma, species near xus, new spe-
cies, etc., can also be entered and retrieved. The system is
flexible enough to handle subgenera and subspecies as well.

(GEOGRAPHY

Emerson and Ross (1965) and Levi (1966) have commented
on the importance of locality data in collections of inverte-
brates, and other authors have cited one or more methods of re-
cording geographic data, including distance and direction from
a known point to a locality (10 mi. N., 4 mi. E) (Riemer, 1954;
Hutchison, 1964), use of township, section, and range (Ax-
tell, 1965), legal description (Wheeler, 1965), and so on. In
studies on marine animals, latitude and longitude ( Axtell,
1965; Steward, 1965) or Marsden Square are commonly used.
Ail of these methods have specific applications; none are used
exclusively by all taxonomists.

For these reasons we have had to develop our own geo-
graphical code, called the Global Reference Code, designed by
Reginald Creighton, Anthony Piacesci and Dick King of the
Smithsonian Information Systems Center when it became ap-
parent that existing methods of storing retrieving geographic
data had too many limitations. Creighton and his colleagues
are preparing a paper on development of the Code.

The Global Reference Code is a hierarchical system in that
several levels, from the general, such as Pacific Ocean, to spe-
cific, such as Manila Bay are used, with room for four levels of
complexity. Locality data may be entered by latitude and
longitude or by name.

682 Proceedings of the Biological Society of Washington

Latitude and longitude are assigned to specific localities by
hand, and a separate geographic data bank is maintained by the
computer.

The smallest area defined by the GRC is a two-minute square.
All data referring to localities within each two-minute square
can be retrieved and printed out as originally entered.

The following example will demonstrate the flexibility of the
system. Soldier Key is a small island on the eastern edge of
southern Biscayne Bay, Dade County, Miami, Florida. It may
be identified, in different collections within the museum, as
(a) Soldier Key, (b) a small isolated key 10 miles south of Key
Biscayne, Miami, (c) a key north of Elliot Key, Biscayne Bay,
or (d) it may have been identified originally by its coordinates
or by compass bearing and range from a point. If information
on species found at Soldier Key is required, the computer will
select and print out all of the data given above, as originally
entered.

This has numerous benefits, for we can retrieve data by lati-
tude and longitude, ocean, county, state, drainage system,
Marsden Square, or by any of existing methods of recording
locality data.

Eventually, as the geographic data bank develops, routines
can be developed so that it can be searched for specific geo-
graphic data and reenter these into the catalog record auto-
matically.

RESULTS

Initially there was a serious lag between development of
computer programs for data storage and manipulation, as well as
development of formats for geography, so that by the time the
programs were ready a large series of records had accumulated
awaiting entry into the system. We then learned that all of our
entries accumulated in this period had to be redone for a va-
riety of reasons, primarily because the fields for each entry had
not been identified correctly. The beginning and end of each
field must be flagged or marked by the typewriter operator and
the flag, an asterisk or an exclamation point, must appear on the
tape, or, as far as the computer is concerned, the fields cannot
be identified and the record is invalid.

Natural history collection symposium 683

Although we believed we had designed the system so that we
could eventually phase out the ledger catalog, i.e., it would be
replaced by a card file in numerical order, we quickly learned
that a ledger or equivalent, in the form of a work sheet, was
required in order to prepare and arrange the data. We still
use the ledger and we have dropped, at least temporarily, the
numerical card file.

Now that the geographic data bank and the Global Refer-
ence Code has been developed and is functional, we should be
able to replace the geographic card in our cataloging operation.

I have noted above that we learned that verification and as-
sembly of data by a cataloger, in preparation for the actual
typing operation, required the use of a ledger catalog or data
entry sheet of some sort. From an operational point of view it
was much simpler for the trained cataloger, a data specialist,
to work with the specimens in an area not necessarily adjacent
to the typewriter. Alcoholic collections are too messy for the
basic work to be done at or near the typewriter. Initially, we
used the cataloger not only as a data specialist but also as a
typewriter operator. We have learned from our first year of op-
erations that the cataloger can be reserved for the data-verifica-
tion aspects of the operation, and that the data can be entered
by a clerk-typist. Further, a clerk-typist can enter data far
faster than any one cataloger can prepare it.

As we recataloged the stomatopods, data for the types was
entered along with data for all of the other specimens in the
collection. In working with the preliminary printout, the Work-
In-Progress listing, of the data from the types, it became ap-
parent that it would be relatively simple to reformat the data
and generate a type-catalog by the computer.

A program has been developed and the printout has been re-
quested. We expect to obtain in one printout an alphabetized
list of the types, as cataloged under their original names, along
with a list of the current names for those that have changed
since the original description.

Another finding is that our data input is not completely satis-
factory, for we need the capability of including more habitat
data. We had reserved space under “Remarks” for habitat data,
reference to field records, color notes, and so on. In the case of

684 Proceedings of the Biological Society of Washington

many of our specimens, there is relatively little information
available other than locality, date, and collector. Now the
number of records with habitat data is so slight that our in-
quiries can be worded so that only records with information in
the “Remarks” section are printed. We will be working on this.

On the cards themselves, we have reserved space at the bot-
tom to edge-punch the cards with the basic data for invoices;
catalog number, name, and total number of specimens. If the
cards were punched as they were processed, we could even-
tually be in a position to generate invoices of loans from our
cataloged collection by stacking cards, in the order desired, in
the Typetronic reader and let the machine automatically type
out the basic data. Unfortunately, edge punching obliterates
any entries in the “Remarks” section so we have not imple-
mented this as yet.

POSSIBLE EXPERIMENTAL PROGRAMS

During the second part of the project we plan to begin manip-
ulation of the data base built up during the first 1S months of
operation. Interrogations designed to test the capabilities of
the system and to provide information as retrieval costs might
include the following:

(1) List and count all species in genus represented in collec-
tion.

(2) Determine number of species in given collection and state
source.

(3) List all materials identified from accession Z.

(4) What species of Family X occur together in depth range
X-Y in the northwestern Atlantic. Plot distribution pattern
of each species.

(5) What species in genus X are not represented in the col-
lections.

(6) List by accession number material of family Y from the
eastern Pacific not yet identified to species.

(7) List species occurring at island X.

(8) List species collected by ALBATROSS at Sta. X.

(9) List type-species of family X or order Y not represented by
materials in collection.

Natural history collection symposium 685

In addition to working out various queries and testing the
system, I would like to work on several items. I have already
noted that in some cases adequate data, available from field
notes, etc., cannot be entered on the 3 X 5 specimen card. I
would like to develop a station data Directory, similar to our
geographic Directory, in which all data pertaining to a partic-
ular station or collection could be entered and also be availa-
ble for combination with the basic catalog data by the com-
puter.

For example, Dr. Waldo L. Schmitt kept extensive field notes
which relate directly to many specimens now in the crustacean
collection. If we could enter all of this by station number and
have the computer add this data to each printout record, we
could greatly increase the amount of information available on
each specimen. For the purposes of management of our collec-
tion, this information is not neccessarily needed in our species
file. Its value for research purposes is obvious.

Similarly, data from Smithsonian expeditions and other ex-
peditions as well might be stored and tied to the specimen rec-
ord by the station number.

We also plan to develop a basic catalog card for parasites
and commensals from our crustacean holdings which will
emphasize host and ecological data.

During the coming year we plan to investigate various ap-
plications of the computer and the computer-based specimen
data bank to our routine operations. The possibilities are un-
limited. We should be able to generate invoices giving com-
plete specimen data, lists of holdings for visitors, lists of hold-
ings for exchange purposes, as well as a machine-generated
catalog in tabular form which could be bound and retained as
a permanent record. Entry of present unidentified material
(identified to family or genus level, but not species level), would
enhance our abilities to make our collections and these data
available to visitors.

Development of off-campus storage facilities for inactive
collections, in my opinion, is dependent on development of
banks of data associated with those collections; removal of col-
lections to off-campus facilities without documentation of the
material would be tantamount to destruction of the material.

686 Proceedings of the Biological Society of Washington

I would also like to see the museum develop methods for
computer storage of records from the literature as well as data
associated with specimens in our collection.

LITERATURE CITED

AXTELL, RALPH W. 1965. More on locality data and its presentation.
Systematic Zoology, 14 (1): 64-66.

BLACKWELDER, R. E. 1963. Classification of the animal kingdom. South-
ern Illinois University Press. Carbondale, Hlinois.

Buuuis, Harvey R., Jr., AND RicHArp B. Ror. 1967. A bionumeric code
application in handling complex and massive faunal data. Sys-
tematic Zoology, 16 (1): 52-55.

EMERSON, WILLIAM K., AND ARNOLD Ross. 1965. Invertebrate collections:
Trash or Treasure. Curator, 8 (4): 333-346.
Levi, Herpert W. 1966. The care of alcoholic collections of small in-
vertebrates. Systematic Zoology, 15 (3): 183-188.
Hutcuison, Vicror H. 1964. Distance and direction in locality data.
Systematic Zoology, 13 (3): 156-157.

RieEMeER, WiLtiAM J. 1954. Formulation of locality data. Systematic
Zoology, 3 (3): 138-140.

Sgurres, DonaLD F, 1966. Data processing and museum collections: a
problem for the present. Curator, 9 (3): 216-227.

STEWARD, CHARLES C. 1965. More on latitude and longitude. Systematic
Zoology, 14 (4): 343-344.

WHEELER, GEORGE C. 1965. Locality by legal description. Systematic
Zoology, 14 (1): 66-68.

THE HERBARIUM: PAST, PRESENT, AND FUTURE!

By STANWYN G. SHETLER”
Smithsonian Institution, Washington, D. C.

The herbarium as an institution dates back more than four
centuries, but the origins of plant collecting for medical and
proto-scientific purposes trace back even further to the time of
the medieval herbalists. To some of today’s biologists the her-
barium is an anachronism in the modern scientific world, and
their voice of reproach has seemed to grow ever more deafen-
ing, especially to the ears of the curator. In response have come
some eloquent defenses of the herbarium and its historical
significance to science and human welfare (e.g., Beaman et al.
1965, Cronquist 1966). Notwithstanding frequent optimistic
predictions, however, today the herbarium faces critical chal-
lenges to its future existence. Quantitative gains too often are
being mistaken for progress and good health, while in fact the
physical facilities, staff, operational procedures, collection strat-
egy, and intellectual raison d’étre of the herbarium are not

1 This paper, given at the symposium under the title “The Future of the Herbar-
ium,” has been expanded greatly in the statistical portions dealing with the past and
present of the herbarium. “A herbarium,” as Lawrence (1951) defines it, “is a col-
jection of plant specimens that usually have been dried and pressed, are arranged
in the sequence of an accepted classification [can be purely alphabetical], and are
available for reference or other scientific study.’’ Cronquist (1966) would add that
“a herbarium can be a very useful teaching aid or an absorbing hobby. .. .” It is
beyond the intent of this paper to review the extensive and widely scattered literature
on the history, philosophy, apologetics, and methodology of herbaria.

2JT am deeply indebted to Elaine R. Shetler and Nancy L. Howard for help in ab-
stracting, keypunching, and compiling the statistics, and to James J. Crockett and
Shigeko I. Rakosi, Smithsonian Information Systems Division, for taking care of the
computer programming and processing. Dr. John H. Beaman, Curator of the Beal-
Darlington Herbarium of Vascular Plants, has kindly cooperated in supplying infor-
mation on the Michigan State University Herbarium. Acknowledgment is due also
to Mildred J. Davenport and Betty Scott for assistance on the manuscript. Financial
support was provided by a grant (Sg0621054/C-1) from the Smithsonian Research
Awards Program.

56—Proc. Biot. Soc. WaAsH., Vou. 82, 1969 (687 )

688 Proceedings of the Biological Society of Washington

keeping pace with the times. Future prospects are being fore-
cast on the basis of the past or present significance of the her-
barium without due regard for its changing role. Conse-
quently, the prognoses are at best too optimistic and at worst
delusive or irrelevant.

In many respects, to be sure, the ills of the modern herbarium
are only symptomatic of the greater malaise that besets the
whole of classical botany and indeed biology (Bonner 1963,
Laetsch 1963, Shetler 1963, Smith 1964, Engledow 1968). On
the one hand, the traditional disciplinary approach to biology,
which has tended to partition it into botany and zoology and
then into kinds of plants and animals, is giving way to the
levels-of-organization approach, which is topical and cuts
across the classic groupings of organisms. On the other hand,
descriptive biology at the higher levels of organization is be-
coming unfashionable and is being supplanted aggressively in
curriculums and graduate research programs by descriptive and
experimental molecular biology. Fortunately there are scien-
tists who understand the importance of descriptive biology at
the higher levels and can counteract the trend to supplant
rather than complement such biology with molecular biology
(e.g., Mayr 1968).

Faced with the crisis in classical biology, the museum, that
citadel of descriptive biology of which the herbarium is but a
special case, is remarkably healthy and viable today even in
many universities. In some instances, in fact, other, seemingly
more favored scientific facilities have taken second place to the
museum. There is always the overwhelming physical reality of
a large collection of specimens that cannot be ignored easily,
although it is precisely this attribute that increasingly has be-
come a negative factor whenever the future of a museum is at
stake.

Whether the omens for the future seem favorable or unfavor-
able the time has arrived for curators to take a realistic look at
the current plight of the herbarium and to make some frank
assessments of future needs and prospects. This must be done
at the risk of a misuse of findings among our critics. I hope
that this paper will provoke more dialogue among herbarium
curators, administrators, and plant systematists at large con-

Natural history collection symposium 689

cerning the future of the herbarium and thereby lead to a
more exhaustive study of the question than I am able to offer.

SourRcE OF STATISTICS

The herbarium fraternity, thanks to the pioneer efforts of
Professor Lanjouw (see “Introduction,” Lanjouw and Stafleu
1959), has been polling itself for many years concerning her-
barium resources. This work has been carried out under the
auspices of the International Bureau for Plant Taxonomy and
Nomenclature of the International Association for Plant Tax-
onomy (IAPT), with headquarters in Utrecht, Netherlands.
The results have been published in Index Herbariorum, Part I:
The Herbaria of the World (hereinafter abbreviated I.H.),
now in its fifth edition (Lanjouw and Stafleu 1964), with a
sixth due this year. This compilation, though it has obvious
shortcomings, is invaluable and unique: no other group of biol-
ogists, to my knowledge, has such a concise, worldwide digest
of its research collections. Other, complementary reference
guides published by the IAPT are an index to plant collectors
(Lanjouw and Stafleu 1954, 1957), an index to institutional
wood collections (Stern 1967 ), a directory of plant taxonomists
(De Roon 1958), and a directory of botanical gardens (How-
ard et al. 1963). Together these reference works constitute a
gold mine of information that could be exploited more fully if
they were computerized for easy permutation and comparison
of the data. Perhaps this will be done in future editions.

My analysis is based largely on statistics abstracted from the
most recent edition (5th) of I. H. and permuted by computer.
Dated 1964, this edition is effective only through 1963, thereby
providing a 5-year supplement (1959-63 inclusive) to the
fourth edition (Lanjouw and Stafleu 1959). Ten data fields
were formatted on an IBM card, and a card was keypunched
for each herbarium treated in the text. Geographic data were
supplemented from a world atlas. The ten fields are: (1) of-
ficial herbarium abbreviation (e.g., US for U.S. National Her-
barium, Smithsonian Institution ),* (2) city, (3) state or prov-

3 In the discussion that follows, I sometimes have given only the standard abbrevia-
tion of a herbarium in lieu of the full name, so that the general reader will be spared

meaningless details. With the abbreviation, taxonomists who are interested can look
up the specifics in I.H., which they usually have close at hand.

690 Proceedings of the Biological Society of Washington

ince (only for Australia, Brazil, Canada, China, Great Britain,
India, Mexico, USA, USSR), (4) country, (5) continent or re-
gion, (6) year of founding, (7) number of specimens, (8) num-
ber of staff, (9) organizational status (university, government,
private), and (10) type of plants (phanerogams, cryptogams,
general ).

With the aid of a Honeywell 1250 computer, a directory
(Shetler et al. 1968) was produced indexing the herbaria al-
phabetically by: (1) abbreviation; (2) city; (3) country and
state or province within country; (4) continent, country within
continent, and state or province within country; (5) organiza-
tional status and country within status; and (6) type of plants
and country within type. Also, the herbaria were ordered by
(7) year of founding, (8) size of collection, and (9) size of
staff. In the process of indexing, certain statistics were com-
puted by machine, and still other statistics have been computed
manually from the printout for the purposes of this paper.

Of the 1,188 herbaria listed in I.H. (1964) at least by name
and abbreviation, only 941 are actually treated or mentioned in
the text, and 8 of these (BM-SL, G-DC, ND-G, SAM, SARF,
STE-VB, TM, TRV) are incorporated with other herbaria and
do not have separate statistics.1 This leaves 933 herbaria for
which at least some data are given. Unless otherwise indicated,
all statistics and comparisons are based on an analysis of data
provided for these 933 herbaria.

Index Herbariorum is intended to cover only public, institu-
tional herbaria. Collections in the hands of private individuals
are not considered part of the public domain of science and are
not assigned standard abbreviations (mark of official recogni-
tion) nor included in I.H. The hundreds, probably thousands,
of private herbaria in the world are usually small, seldom ex-
ceeding a few hundred or thousand specimens. In at least
one case, however, a private collection is known to number
about 150,000 specimens, a not insignificant herbarium. Of

4 The alphabetical index to herbarium abbreviations registers 247 herbaria omitted
from the text, all but one (GUA) being small British institutional herbaria taken from
Kent’s book (1957) and listed in I.H. to provide conveniently their official abbrevia-
tions. Twelve of the 941 herbaria treated or mentioned in the text are not included
in the index: BM-SL, CHIS, CHISA, G-DC, KL, KLA, KLU, KRA, ND-G, SARF,
TENN, TM.

Natural history collection symposium 691

course many institutional herbaria began as private collections.

Even as a register of public herbaria, I.H. still falls short of
completeness after 30 years of data-collecting and updating.
Of the 167 herbaria reporting for the first time in the 5th edi-
tion, which represent about 18% of the 933 herbaria treated,
only 13 report a founding date in the years (1959-63) since the
4th edition appeared. Thus even in the latest edition of I.H.
over 16 percent of the main entries (92 percent of new entries )
are entries that should have appeared already in the 4th edi-
tion if not before. The real total of public institutional herbaria
has not been approached. We know in the case of Great Brit-
ain, thanks to Kent's book (1957), that almost five times as
many institutional herbaria have escaped full treatment in I.H.
as have been treated (246:50). Every country, no doubt, has
its own small, unnoticed herbaria in municipal, county, and
state or provincial museums, schools, and parks. The United
States, for example, has many, often quite valuable though lo-
cal herbaria in national parks. For the most part, these obscure
herbaria, which so far have either failed to respond to question-
naires or have escaped the notice of the compilers of I.H., are
small and inactive with respect to the national and international
commerce of plant taxonomy. It is likely that the number of
scientifically important herbaria in the world is about 1,000, i-e.,
approximately the number now treated in I.H. Doubtlessly
some important herbaria, particularly in the USSR, China, and
Southeast Asia, have not yet reported, but at the same time
some of the tiny herbaria already treated in I.H. are relatively
unimportant to the pursuit of systematics. Having said this, I
hasten to add that in a real, if relative, sense all herbaria are
scientifically important. It is to be hoped that eventually [.H.
can be a complete worldwide register of institutional herbaria.
If the 5:1 ratio of unreported to reported herbaria of
Great Britain were to hold throughout the world, then there
could be as many as 5,000 institutional herbaria. If Kent’s data
are a sate guide to the size of the smaller, unreported herbaria
of the world, then such herbaria have anywhere from 200 to
75,000 specimens and average almost 5,000 specimens /herbar-
ium. At this rate, 1-1.5 million specimens should be added to

692 Proceedings of the Biological Society of Washington

the I.H. figure just for Great Britain, and on a worldwide scale
this could mean an additional 18-20 million specimens.

The problem of missing data is bothersome because many
herbaria did not report complete information. Number of speci-
mens was reported by 78 precent of the 933 herbaria and year
of founding by 79 percent, while 85 percent listed the names of
one or more staff members. Perhaps some of the 15 percent not
listing staff in fact do not have any staff. Except where other-
wise indicated, the statistics of this paper are based on the her-
baria actually reporting and are not extrapolated to account for
all 933 herbaria treated in I.H., to say nothing of the 247 men-
tioned but not treated in I.H. or of any estimated world total of
herbaria. It should be kept in mind, therefore, that in reality
the figures would be higher, perhaps much higher, in all cate-
gories if data were available for all public herbaria. The bias
of missing data probably affects the statistics for most countries
about the same, but there are some notable exceptions. The
herbarium resources of mainland China cannot be assessed real-
istically because 73 percent of the entries in [.H. for Chinese
herbaria (excluding Taiwan) do not include number of speci-
mens or the names of staff. For the USSR, only 58 percent of
the included herbaria report staff and only 60 percent report
number of specimens. French herbaria report number of speci-
mens in even fewer cases (57 percent). Some of the smaller
countries have not reported any staff or specimen totals.°

Concerning the reliability of the data in I.H., the questions,
When is a herbarium actually founded?, What is a specimen?,
and, Who is a staff member?, naturally arise.

Establishing the founding date of a herbarium can be a quite
subjective matter. The U.S. National Herbarium,® for example,

5 Countries not reporting any staff (total number of herbaria in parentheses): Brit-
ish Honduras (1), Ecuador (2), Greenland (1), Lebanon (1), Nicaragua (1),
Paraguay (1), Ryukyu Islands (1), and Seychelles (1); countries not reporting any
specimen totals: Azores (1), British Solomon Islands (1), Ecuador (2), Greenland
(1), Korea (2), Nicaragua (1), Paraguay (1), Ryukyu Islands (1), and Tunisia
(1).

6 The U. S. National Herbarium, as the Smithsonian’s plant collection has long
been designated in the international taxonomic fraternity, is administered by the
Department of Botany of the Institution’s National Museum of Natural History.
Technically speaking, therefore, ““U. S$. National Herbarium’? is a term of conve-
nience for the collections themselves and not an official organization with a staff and

administrative status. For practical purposes, however, it can be so regarded in
many contexts.

Natural history collection symposium 693

gives a founding date of 1868 in I.H. This was the year when a
Smithsonian herbarium was organized in Washington, D. C.,
under the care of the U. S. Department of Agriculture, but
shortly after its founding in 1546 the Smithsonian Institution
had already come into possession of plant collections made
under federal auspices as early as 1840 (Stern 1966). It was not
until 1894, however, that the U. S. National Herbarium was of-
ficially established at the Smithsonian. A further example is the
herbarium of the Komarov Botanical Institute in Leningrad,
said in I.H. to have been founded in 1823, but which actually
was an outgrowth of collections started almost at the inception
in 1714 of the forerunner medical garden (cf. Shetler 1967,
Lipschitz and Vassilezenko 1968). In I.H7., the founding dates
of the medical garden and herbarium are distinguished from
each other. Even when, as in this case, the distinction is made
in I.H. between the founding dates of the herbarium and its
mother institution, choice of starting point may be entirely sub-
jecuive:

The overwhelming majority of specimens reported in I.H. are
of the conventional herbarium type, but it is clear that other
types of specimens (fossils; wood samples; fossil or wood thin
sections; pollen, spore, and other anatomical microscope slides )
frequently are included in the totals. Cryptogamic specimens
are especially problematic. There is no uniform way of count-
ing them; yet generally they are not tallied separately in I.H.
One must assume that the confounding effects of cryptogamic
and other kinds of specimens are spread over all herbaria.

To judge by the few herbaria giving exact figures for totals,
one would conclude that only about 3 percent of the world’s
herbaria actually maintain precise counts of specimens held.
Furthermore, it is not possible to know in any given case how
many of the specimens of the total are mounted as opposed to
unmounted or available for consultation as opposed to being in
storage and unavailable.

The criteria for reporting staff obviously varied from one her-
barium to another. In general, only professional curatorial-re-
search staff are listed, but some universities and research in-
stitutes have reported whole faculties or groups of faculties, so
inflating their actual staffs that it is impossible to know how

694 Proceedings of the Biological Society of Washington

many persons play an active role in the herbarium. Other in-
stitutions have included directors or administrators who have
nothing to do with the herbarium and in fact may not even be
botanists, while still others have included technical staff such
as preparators. In the future, herbaria should be encouraged to
report total number of technical and clerical staff, without
names, and to distinguish, as some herbaria have already done,
between active curators on the one hand and associated re-
searchers and emeritus or honorary curators on the other hand,
so that an accurate picture of the world’s professional man-
power devoted to the maintenance of herbarium collections can
be ascertained.

Lacking any sound basis for consistently distinguishing be-
tween different kinds of staff in I.H., I have simply counted all
persons listed for each herbarium. If we can assume that the
excesses of one are cancelled by the deficiencies of another,
then we can assume that the total figures yield a reasonably fair
report of professional curatorial manpower in the world’s her-
baria. Given this rough level of manpower estimation, I have
ignored cases of duplication. About 2 percent of the names ap-
pear twice, but often it is not possible to know whether the
curator was holding two positions or had moved to another
herbarium too recently for his name to have been removed from
the roster of the first herbarium.

Notwithstanding its limitations and shortcomings, Index
Herbariorum provides an excellent statistical abstract of the
world’s herbarium resources for which the compilers must be
given full credit. Even though the data in specific cases may
be suspect, this should not invalidate collective statistics and
comparisons unless there is evidence of systematic bias. I hope
that my paper will have the positive effect of stimulating cura-
tors to help in correcting and refining the data in future edi-
tions of I.H. where necessary.

To supplement the data of J.H. and make my analysis more
vivid and contemporary, I have selected two American herbaria
for brief case study. As an example of a university herbar-
ium, I take the Michigan State University (MSU) Herbarium.
A recent symposium, held about the time when the 5th edition
of I.H. was issued, focussed attention on the MSU Herbarium

Natural history collection symposium 695

and the problems of university herbaria in general. This herbar-
ium, with its approximately 200,000 specimens, characterizes
active university herbaria of small to moderate size. The U. S.
National Herbarium will serve as my example of a large non-
university herbarium. For statistics on the National Herbarium
I have drawn freely from the excellent status reports of the
Smithsonian’s Department of Botany that were prepared for
internal purposes recently by Stern (1966) and Hale (1967).

GrowtH Or Worwp’s HERBARIUM RESOURCES

All of the tables (1-16), compiled from the 5th edition of
I.H. (1964), are placed at the end of the paper.

Chronology of Herbarium Founding

The first institutional herbarium was founded about 425
years ago in 1545 at the University of Padua in Italy’ (see
Tables 5-12). Only 12 more herbaria were to be founded dur-
ing the next 200 years, including four others in the 16th century
and four in the 17th century. Among the latter were two,
formed in 1635, which today are among the world’s most re-
nowned herbaria, namely, the phanerogamic (P) and crypto-
gamic (PC) herbaria of the Muséum National d'Histoire Nat-
urelle in Paris. The first half of the 18th century saw only four
herbaria established, but the second half brought a minor burst
of 32 foundings. Several of the great herbaria of Europe and
the British Isles took origin during this period. Thus herbar-
ium formation did not begin in earnest until about 1750. The
year 1753, when Linnaeus published his revolutionary Species
Plantarum, was only the second in history in which two her-
baria were founded (British Museum, London; University of
Vienna, Austria), and the decade of the 1750s was the first in
history in which more than two (five) herbaria were formed.
Henceforth, the number of herbaria formed per decade, plotted
in Fig. 1 for every decade from 1750-59 to 1950-59, began to
rise, only twice dipping below the level of the 1750s. Of the 21
decades from the 1540s to the 1740s, by contrast, there were 10

7 The founding date for the university herbarium at Pisa, Italy, is given in I.H. as
“hefore 1850,”’ but the botanic garden of the university, which may have maintained
a dried plant collection early in its history, is said to have originated in 1543. The
analysis deals, of course, only with herbaria in existence.

696 Proceedings of the Biological Society of Washington

in which no herbarium was founded, 9 in which only a single
herbarium was formed, and 2 (1560s, 1630s) when two her-
baria were founded.

Prior to 1750, only one herbarium (Mauritius, 1737) had
been formed outside the continent of Europe. Several herbaria
came into being in the British Isles during the next 50 years,
and the first herbarium of the New World was founded in 1772
at Winston-Salem, North Carolina, in the United States. A year
later the second and only other New World herbarium to be
founded prior to the 19th century was formed at Charleston,
South Carolina, where the collections of Stephen Elliott,
pioneer botanist of the Carolinas, have been kept. Neither of
these herbaria ever advanced far. The first principal herbarium
of the United States, though actually the fourth to be founded
in this country, was organized in 1812 at the Academy of Nat-
ural sciences of Philadelphia (given as first American herbar-
ium by Jones and Meadows 1948). In Asia, the first herbarium
was established in 1793 at Calcutta, India, and a second was
not formed until almost 25 years later. The first herbaria of
South America and Africa were not formed until the 1800s, in
1808 ( Rio de Janeiro, Brazil, RB) and 1855 (Cape Town, South
Africa, SAM), respectively.

Up to the year 1800, i.e., for over 250 years, only 45 herbaria
had been founded, 94 percent of these in Europe and the Brit-
ish Isles. During the next 50 years (1800-49 ) the pace of found-
ing quickened markedly, and 76 herbaria, about 1.5/year, were
formed. Well over half of these were formed in Europe and
the British Isles, but a dozen were founded in North America
and a handful elsewhere. The rise and spread of the herbarium
as a scientific institution had really begun. Thus, while only 13
herbaria had been established in the entire world prior to 1750,
the British Isles and North America each had about this many
by 1850. By contrast, however, Asia, the Australasian-Pacific
Island region, and South America were not to achieve about a
dozen herbaria each until 1900, more than 350 years after the
very first herbarium was organized, and Africa could not claim
this milestone until the first decade of the present century had
passed.

During the 1800s, 270 (37 percent ) of the present 933 herbaria

Natural history collection symposium 697

were established, at an average rate of 2.7/year. By the late
1830s one or more herbaria were being created virtually every
year, and, according to the official record (I.H.), over the 125-
year period from 1839 to 1963 there have been only 5 years
(1841, 1843, 1851, 1866, 1961), including only one in the present
century, when new herbaria have not been founded. The real
explosion in herbarium building has come during the present
century. Thus far (1900-63) 420 herbaria, 57 percent of the
total number, have been founded, averaging 6.6/year, which is
more than twice the rate for the 19th century and exactly triple
the overall rate (2.2/year) for the whole 419-year period
(1545-1963 ). Of these 419 years there have been 239, of which
234 occurred prior to 1839, when not a single new herbarium
was formed.

The golden age of herbarium-founding began about the mid-
dle of the 19th century and lasted for about 100 years (Fig. 1).
The 1850s witnessed a sharp upswing to 30 in the number of
herbaria founded per decade, and from there the general trend
was upward until the 1920s when 91 herbaria, the all-time high
for a single decade, were formed. The peak year was reached
in 1890 when 20 herbaria were founded. There have been only
13 other years in history, all in the 20th century, when 10 or
more herbaria were founded in a single year; these vintage
years, in order of decreasing number of herbaria formed, have
been: 1930 (15 herbaria); 1920 (14); 1935, 1947 (13); 1918,
1925, 1946 (12); 1900, 1922, 1923, 1924, 1950 (11); and 1932
(10). Since the 1920s, the founding of new herbaria has de-
clined sharply.

The 100-year herbarium boom has coincided roughly with a
similar golden age of exploration and description in plant sys-
tematics and biology generally, which was initiated by the
great pioneer biologists of the late 18th and 19th centuries and,
it appears, is now drawing to a close in mid-20th century. A
pivotal factor in the United States, both in the flourishing of ex-
plorative-descriptive biology and in the rise of the herbarium,
was the passage by the U. S. Congress of the first (1862) and
second (1890) Morrill acts (sponsored by Representative Justin
Smith Morrill), providing for the establishment and support of
land-grant colleges to promote, among other studies, the agri-

698 Proceedings of the Biological Society of Washington

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cultural sciences. Some of the most outstanding herbaria in the
United States today are to be found in land-grant institutions
tracing their starts to this legislation. Although the influence
and support of the land-grant acts continue, reinforced by
additional legislation as recently as 1960, the impetus of the
agricultural college, which so profoundly affected the course of
herbarium development in America, was largely dissipated
after the first few decades of the present century. This develop-
ment in America must be regarded as having influenced the
worldwide trend as well, given that the USA can claim nearly
a quarter of the world’s herbaria today.

Herbarium-founding has by no means stopped today, despite
the sharply declining rate since the 1920s. If, as so frequently is
done, one looks only at the present and ignores the historical
trend, then herbaria appear to be growing and spreading more
rapidly than ever, which actually is true in some parts of the
world where the last five or ten years have chalked up a larger
roster of foundings than the first 200-350 years (cf. Tables 5-12
and next section). Worldwide, about 300 years passed before
the first 100 herbaria were formed, but the last 100 of the 933
herbaria have been founded in just over 17 years (Table
14)! During the most recent 5-year period for which
statistics are available, 13 herbaria have been founded—as

Natural history collection symposium 699

many as were founded during the entire first 200 years (to
1750). Obviously the herbarium is far from being a dead insti-
tution. Yet it also must be noted that the annual rate of found-
ings has decreased from 6.6 herbaria/year during the first 63
years of this century to 4.2 during the last 10 years and 2.6 dur-
ing the last 5 years of this period.

Geography of Herbaria

The world’s herbarium resources are analyzed by continent
or region in Tables 1 and 2 and by country (top 22 countries
only ) in Tables 3 and 4.

Today's 933 herbaria are located in 104 countries, averaging
about 9 herbaria, or 1 percent of the total, per country. In fact,
however, only 22 countries have the average 9 herbaria or
more. Of the 104 countries, 37 have only one herbarium, 77
have five or less, and 95 have twenty-five or less. The top five
countries and their numbers of herbaria are: USA (244),
Great Britain (50), Canada (48),* USSR (43), and France
(42). Only the first place of the United States is clearly
established, and this is true whether the criterion is number of
herbaria, number of specimens, or number of staff. The ranks
of the other countries change when the latter two criteria are
used; Canada drops to 13th place when ranked by number of
specimens but only to 5th place by number of staff, while
France places 2nd and Great Britain 3rd by both of these stan-
dards. The rank of the USSR cannot be determined confidently
with the available statistics in [.H. because of the relatively
large number of herbaria not reporting full data; quite possibly
the USSR ranks next to the USA. Likewise, in reality China
might rank among the top five, but the highly incomplete data
place her out of the running.

The 10 leading countries have 63 percent of the world’s her-
baria; none of the other 94 countries has as many as 20 herbaria.
In the United States, every state has at least one herbarium, and

8 On behalf of the Systematics and Phytogeography Section of the Canadian Botan-
ical Association, W. K. W. Baldwin of the National Herbarium of Canada (CAN),
Ottawa, has been making a special study of Canadian herbaria during the past few
years in an effort to provide better data for the forthcoming 6th edition of Index Her-
bariorum, His progress reports, which have been distributed but not published, reveal
that there are more than 60 herbaria in Canada at the present time. I have not at-
tempted to incorporate his incomplete results here.

700 Proceedings of the Biological Society of Washington

California leads with 27. Half of the USA’s 244 herbaria are
found in California and six other states: Texas, 22; Michigan,
19; Massachusetts, 17; Pennsylvania, 14; New York, 12; and
Ohio, 10. If the continents of Europe and North America are
treated broadly, as follows, then each is seen to have about a
third of the world’s herbaria: Europe + British Isles = 327 her-
baria; North America (including Mexico and Central America )
+ West Indies = 315 herbaria. Asia places a distant third
place with 114 herbaria or, including the Australasian-Pacific
Island region, with 158 herbaria.

The 933 herbaria of the world are found in 669 cities, averag-
ing 1.4/city. Thus, inefficient and wasteful as it may be to
maintain two or more facilities and collections within the same
city, such duplication has been common practice. Historical
precedent or petty institutional sovereignty and politics too
often seem to outweigh the scientific logic and simple econom-
ics of consolidation. Ironically, moreover, it seems to be a uni-
versal fiscal principle of bureaucracies, at least governmental
ones, that two small units often can command greater total sup-
port than one large, consolidated unit.

As mentioned earlier, the herbarium was an almost exclu-
sively European institution for more than 200 years. Then slowly
it was transplanted to foreign soil by European naturalist ex-
plorers turned settlers, and over the years the geographic focus
of active herbarium-founding has tended to shift more or less in
phase with the shifting thrust of European and, eventually,
North American exploration and colonization. An indigenous
tradition did not take hold in North America until well into the
19th century nor in Asia and Australasia until as late as the
early 20th century in some parts. Only in quite recent years has
the herbarium become truly indigenous in Africa and South
America.

This shifting focus of activity can be documented statisti-
cally. Table 14 compares the geographical distribution of the
first 100 herbaria founded with the geographical distribution
of the last 100 founded, i.e., counting back from 1963, the most
recent year for which I.H. gives data. The figures, which can
be read directly as percentages, speak for themselves.

During the recent 25-year period from 1939 to 1963, in-

Natural history collection symposium 701

clusive, 150 herbaria were founded, and North America (includ-
ing Mexico but excluding Central America and the West In-
dies) led with 42, followed by Africa with 28, Europe with 26,
South America with 21, and Asia with 19. If only the last 10
years of this period are considered, then Africa noses out North
America by one herbarium (9:8). Herbarium-founding has
tapered off greatly in Europe, where on the continent not a
single new herbarium was formed during one recent 5-year
period (1954-59), and several countries apparently have not
founded a new herbarium this century; nevertheless, it is quite
remarkable that any herbaria at all are still being started here,
as at Aarhus, Denmark, in 1963. This speaks well for the con-
tinuing vitality of descriptive botany in Europe. In North Amer-
ica, the focus of active herbarium-founding is shifting from the
United States to Canada. A surge of herbarium-building in
Mexico comparable to that taking place recently in Canada has
yet to begin.

The countries of Africa and South America probably never
will experience a herbarium boom to equal that experienced in
Europe or North America. For one thing, Europeans and North
Americans continue to do a large portion of the tropical collect-
ing and research and, therefore, to carry most of the spoils of
exploration back to their home institutions. Furthermore, the
present rapid evolution of biology away from the descriptive
stages, the accelerating pace of the race to conclude the botan-
ical exploration of the earth, and the growing worldwide con-
cern about overpopulation and its destruction of our natural en-
vironment—all seem to be foreclosing on any new herbarium
boom of the scale witnessed in north temperate regions by the
last 100 years. The coming of rapid, easy means of transporta-
tion during the present century has greatly stimulated and fa-
cilitated worldwide exploration. Seemingly, however, modern
means of travel have served mainly to aggrandize the long-
established herbaria and have stifled rather than stimulated the
creation of new herbaria, because in a jet age the remotest parts
of the world are but a research grant away from any would-be
collector's home base. At the same time, representing a grow-
ing, unpredictable counterforce, which in many countries
(e.g., in Africa) already has curbed expiorations by foreigners

702 Proceedings of the Biological Society of Washington

and spurred much new, indigenous herbarium-building, is the
rising tide of nationalism that generates demands for national
science and scientific institutions.

Obviously, many factors may govern the development of
herbaria within a country. Size of home territorial area may
be least among them, witness Great Britain and the countries of
Europe. By contrast, the huge size and floristic diversity of the
Soviet Union have been major factors in keeping most Russian
botanists at home through the years, while at the same time
this size and diversity have enabled them to amass large and rich
collections (Shetler 1967). The impact of the land-grant legisla-
tion in the United States, discussed above, demonstrates the
obvious point that the development of herbaria within a coun-
try is closely dependent on the general level of educational,
scientific, and economic development of the country as a whole.
A country that does not have mature scientific traditions and
institutions also will not have well-developed herbaria nor the
scientific and educational foundations to support them. Every
country goes through a predictable golden age of its own with
respect to the formation of indigenous herbaria, and this curve
is a minor reflection of the country’s curve of overall develop-
ment. Political considerations, especially as they have governed
the national and international movements of botanical collec-
tors and their specimens, have often limited the character and
scale of herbarium-building in a country at least for a time. The
prime modern example of this is to be found in China.

Organizational Status of Herbaria

About 59 percent of the world’s herbaria are university-affili-
ated, 34 percent government-affiliated, and 7 percent indepen-
dent. This classification does not indicate necessarily the
source of funds. In the United States, for instance, virtually no
public herbarium operates entirely on private funds today;
county, state, or federal funds provide at least some support.
The dominance of university herbaria speaks for the impor-
tance traditionally accorded to plant collections in academic bo-
tancial research and education. The rapid increase of herbaria
in North America during the past 100 years has been due in
large part to the rapid increase of state and provincial univer-

Natural history collection symposium 703

sities, many of which have botany departments or botanical
gardens with associated herbaria. The influence of the land-
grant acts in the United States has already been mentioned, and
to this should be added the exemplary, early influence of prestig-
ious schools and teachers. Concerning the development of the
herbarium as an integral element of university botany, it would
be hard to overestimate the profound influence of men like Asa
Gray (1810-88) and Liberty Hyde Bailey (1858-1954) or of
the institutions they served. Since the time of Asa Gray, some
of America’s foremost academic botanists, indeed scientists,
have been herbarium scholars who have made the university
herbarium a primary locus of research and teaching. In the
United States today, Jones and Meadows (1948) point out,

. . almost without exception no first-class university has a
second-class herbarium... .” “Likewise,” they comment, “there
seems to be a very close connection between development and
utilization of the herbarium and the vigor and prestige of a
botanical department.” Chairmen of university botanical de-
partments would do well to savor these observations. Surely
the same kinds of comments about the historic role of academic
herbaria could be made for many countries.

When only the 17 largest herbaria, with 2 million or more
specimens each, are considered, then 53 percent are seen to
be government-affiliated, 35 percent university-affiliated, and
12 percent independent. The relatively higher percentage of
government-affiliated herbaria in this group than among her-
baria at large reflects the fact that government herbaria are
often among the earliest to be founded in a country and they
tend to receive greater and more stable support through the
years than other herbaria, enabling them to grow larger than
others.

Types of Herbaria

Historically, it has not been customary to develop crypto-
gamic and phanerogamic herbaria as separate institutions, al-
though many general herbaria have been organized into differ-
ent laboratories or divisions for different groups of plants. Ac-
cording to available data in I.H., only about 6 percent of the
world’s 933 herbaria are strictly cryptogamic herbaria. Among

704 Proceedings of the Biological Society of Washington

these, however, are some world-famous institutions, including
three with a half-million or more specimens: Laboratoire de
Cryptogamie, Muséum National d Histoire Naturelle, Paris (PC,
1.2 million); Farlow Herbarium of Cryptogamic Botany, Harv-
ard University, Cambridge, Massachusetts (FH, 1 million); and
National Fungus Collection, U. S$. Department of Agriculture,
Beltsville, Maryland (BPI, 675,000, including Smithsonian fun-
gus collections ). One must hasten to add that there are other
large cryptogamic collections (e.g., at Komaroy Botanical In-
stitute, Leningrad, | million specimens) that are not organized
as independent herbaria. In reporting data for future editions
of Index Herbariorum, institutions should attempt to distin-
guish more carefully, the kinds of collections they hold so that a
better picture of the world’s resources by plant groups can be
gained.

Size of Collections

With 724 (78 percent) of the 933 herbaria reporting size of
collection, specimens total about 148 million. If one were to as-
sume that the same average per herbarium (ca. 205,000 speci-
mens ) holds for the 22 percent not reporting size of collection,
then the extrapolated total for the 933. herbaria would be 190
million specimens. Furthermore, if to the 190 million were
added specimens hidden away in national parks and the small
herbaria of municipal, county, and state or provincial museums,
schools, and parks, then surely the world total for institutional
herbaria would reach 200-225 million and possibly as high as
250 million specimens.

Over 131 million, almost 90 percent, of the 148 million speci-
mens are held by the 22 countries with 9 or more herbaria each
(Table 3). With few exceptions the countries having the most
herbaria also have the most specimens, although the ranking is
different. Some of the European countries that have had her-
baria for a very long time rank comparatively much higher in
number of specimens than in number of herbaria (e.g., Czech-
oslovakia ). About 78 million of the 148 million specimens are
concentrated in Europe. This is more than double North
America’s 36 million specimens, and European herbaria also
average more than twice as many specimens per herbarium as
North American herbaria (Table 1).

Natural history collection symposium 705

As a country, the United States of America, with 34 million
based on 86 percent of its herbaria reporting size of collection,
leads the world in total number of specimens. It has about a
quarter of the world’s specimens (23 percent) as well as her-
baria (26 percent) (Table 3). Apparently it is the only country
that has more than 20 million specimens. The nearest competi-
tor, France, has less than half as many specimens (nearly 15
million ); however, this figure is based on only 57 percent of the
herbaria reporting size of collection, so that the real total could
be well over 20 million. Of the 95 countries for which specimen
totals can be compiled from I.H., 48 report 100,000 or less, and
only 3, including the USA, report more than 10 million. Over a
third of the countries have totals between 25,000 and 250,000
specimens. The average for the 95 countries is just under 1.6
million/country, although only 10 percent of the countries have
totals that fall into the range of the average, i.e., 1-2 million.

Compared by average size of herbarium, the USA, with its
160,142 specimens/herbarium, falls far behind other countries.
The top four countries, their herbaria being the only ones to
average more than a half-million specimens each, are: Swit-
zerland (994,286/herbarium), Sweden (965,875/herbarium ),
Czechoslovakia (650,000/herbarium), and France (609,067 /
herbarium). These are countries with long herbarium tradi-
tions where the existing network of herbaria has been stabilized
for some time, and few if any new herbaria are still being
formed.

Forty-five countries have at least one herbarium each with as
many as 100,000 specimens; 25 countries have at least one
herbarium with 500,000 or more specimens; 19 countries have
at least one herbarium with 1 million or more specimens; and
11 countries can claim at least one herbarium of 2 million or
more specimens. Only 6 countries of the world—France, Great
Britain, Italy, Switzerland, USA, USSR—can boast at least one
herbarium of 3 million or more specimens (Table 13).

The statistics in Table 15, based on the 724 herbaria report-
ing size of collection in I.H., give a good indication of the size-
class distribution of the world’s herbaria. As expected, most
herbaria are relatively small, and few are really large. It ap-

706 Proceedings of the Biological Society of Washington

pears that almost half of the world’s herbaria have no more than
25,000 specimens and that almost three-quarters have no more
than 100,000 specimens; about 10 percent have 250,000 or more,
and only about 5 percent have a million or more specimens.

The 39 “big league” herbaria reporting 1 million or more
specimens are listed with appropriate statistics in Table 13 in
order of decreasing size. This list includes 9 herbaria of the
United States, the country with the largest number of herbaria
that have 1 million or more specimens. There are 17 herbaria
with 2 million or more specimens each, and together they have
57 million specimens, more than a third of the total 148 million.
The 10 herbaria with 3 million or more specimens together
have a total of 41 million specimens. Thus it would appear that
25-30 percent of the world’s herbarium specimens are concen-
trated in 1-2 percent of the world’s herbaria, namely, the
world’s very largest herbaria. It is certain that at least some of
the 209 herbaria for which collection size is not given in I.H.
(e.g., herbarium of British Museum in London) belong in
Table 13, but there is no way to take these into account. One
must assume that in relative terms this table gives an accurate
picture of the world’s largest herbaria and their holdings and
staff,

The largest herbarium in the world unquestionably is the
herbarium of the Royal Botanic Gardens at Kew near London,
England, which in 1963 could boast a staggering 6.5 million
specimens. Second place is open to question, however. On the
basis of the data in Table 13, the clear choice is the Muséum
National d Histoire Naturelle in Paris if, ignoring the adminis-
trative separation into two herbaria (P and PC), the 5 million
phanerogamic and 1.2 million cryptogamic specimens are
added together to make a total of 6.2 million. On the basis of
the phanerogamic herbarium alone, the Paris museum may
stand in third place behind the Komaroy Botanical Institute in
Leningrad, where the phanerogamic and cryptogamic collec-
tions, which are administered as one herbarium, total between 5
or 6 million specimens. The Leningrad herbarium has about 1
million cryptogamic specimens, but there is some confusion
concerning the number of phanerogamic specimens, whether 4
or 5 million (see footnote, Table 13). Inasmuch as cryptogams

Natural history collection symposium 707

are included in the Kew and Leningrad totals, it seems only fair
that Paris be compared on the same basis. The herbarium of
the British Museum is not included in Table 13, but already in
1951 it was estimated by Lawrence (p. 231) to have 4 million
specimens. As of 1963, therefore, it might have placed second
or third in size among the world’s herbaria. One of the world’s
largest herbaria (ca. 4 million specimens ) until World War II
was located in Berlin, but it was destroyed in the war.

The largest herbaria of the New World are found in the
United States, and it is a matter of interpretation which places
first, second, and third. According to Table 13, the herbarium
of the New York Botanical Garden and the U. S. National Her-
barium at the Smithsonian Institution, Washington, D. C., were,
with 3 million specimens each, tied for first place in 1963.
Frequently, however, the six herbaria of Harvard University
(A, AMES, ECON, FH, GH, NEBC), Cambridge, Massachu-
setts, are combined when size comparisons are made, and if
this is done Harvard takes the lead, as of 1963, with 3,540,150
specimens. But if this is done for the Harvard herbaria then the
National Fungus Collection at Beltsville, Maryland (just out-
side Washington, D. C.), which includes the Smithsonian’s
mycological specimens, should be considered part of the U. S.
National Herbarium, and the combined total, as of 1963, was
3,675,000 specimens. HH, furthermore, the other herbaria of the
Washington area (LCU, MARY, NA, USFS) are added to this
figure, then the grand total is 4,335,000 specimens. By the same
token, the 294,000 specimens of the Brooklyn Botanic Garden
should be added to the specimens of the New York Botanical
Garden to give a total of 3,294,000 for greater New York City.
In terms of specimens available within the city, therefore, Wash-
ington is first, followed by Cambridge and then New York.

The smallest herbarium on record is located in Siena, Italy,
and had 492 specimens in 1963. It happens also to be the 9th
oldest herbarium in the world, being founded in 1691. This is
the only herbarium reporting less than 500 specimens, although
three others (HNT, SEY, SPH) report just 500.

The general rate of collection growth is difficult if not im-
possible to determine even for a given time period. Clearly the
relative growth rate has been slowing down through the years

708 Proceedings of the Biological Society of Washington

as the bulk of the world’s collections has been increasing stead-
ily. Absolute growth, i.e., in terms of actual number of speci-
mens coming into herbaria, has increased greatly over the past
100 years or more as the number of herbaria and botanical col-
lectors has increased, but there is definite indication that even
absolute growth is on the decline now. Compared to the total
of about 124 million specimens registered in the 1959 edition of
I.H., the total in the 1964 edition is about 24 million higher. The
13 herbaria founded in the period 1959-63, inclusive, report a
total of only 145,000 specimens; obviously, these herbaria do
not account for a significant portion of the 24-million increase.
Between editions of I.H. the U.S. National Herbarium increased
by about 300,000 specimens or 11 percent, as computed on the
1958 base of 2.7 million. If one assumes that herbaria in gen-
eral increased their holdings by about 10 percent during the 5-
year period (i.e., 2 percent/year), then 12-13 million of the
24 million specimens would represent the growth of collections
in previously registered herbaria. This is about 2.5 million
specimens/year, a not unlikely figure for the whole world. The
other 11-12 million of the 24-million-specimen increment prob-
ably are contributed by the more than 150 herbaria reporting
for the first time in the 1964 edition of I.H. even though they
were founded before 1959 and should have been reporting in
1959 or before. Their specimen total does not represent new
growth, except perhaps for about 10 percent of it.

Today, growth relative to the size of existing collections could
be averaging as low as | percent per year among herbaria in
general, meaning an annual worldwide increment to herbaria
of 1.5-2.0 million specimens. Probably the rate lies closer to 2
percent per year, however, because some herbaria are growing
several times this rate (e.g., Michigan State University Herbar-
ium, 5-10 percent/year). During the last five years Canadian
herbaria have been growing at an average rate of more than 6
percent/year (W. K. W. Baldwin correspondence, 1969).
Manpower

Size of professional staff ranges from | to 46, averaging about
4, persons per herbarium and totals 3,158 persons for the 794
herbaria (85 percent of 933) that list one or more staff mem-

Natural history collection symposium 709

bers. The frequency distribution of the 794 herbaria by size of
staff is as follows:

1 staff member 199 herbaria 25 percent
1 or 2 members 395 50
5 or less members 640 81
10 or less members 142, 93
i11—46 members be 7

Only 11 herbaria, listed in Table 16, report 20 or more staff
members (as of 1963). About 75 persons (2.4 percent of 3,158 )
are listed for two jobs in I.H., so that the total number of dif-
ferent individuals is under 3,100 and the average is about 3.9/
herbarium. (Double employment cannot be distinguished
easily from accidental duplication; see “Source of Statistics.” )
Extrapolating with this average, one concludes that the full 933
herbaria are in the care of more than 3,600 individual curators.
If 2-3 percent of the 3,600 serve in two capacities, then the
total number of professional positions occupied is over 3,700.
Distribution of herbarium staff by continent or region is
shown in Table 2. The largest concentration is in Europe (36
percent). North America (25 percent) takes second place, fol-
lowed by South America (11 percent). If the data for Asian
herbaria were more complete, this continent probably would
place third. There are 96 countries out of the total 104 for
which the staff members of at least one herbarium are listed in
I.H, Of the 96 countries, 37 report 5 or fewer staff members,
while 73 report 25 or less; 9 countries report more than 100 staff
members. Only two countries, the United States with 667 and
France with 220, report more than 200 staff members. The
USA has over 21 percent of the world’s herbarium force, based
on these statistics, and France has 7 percent. (By comparison,
the USA has 26 percent of the world’s herbaria and 23 percent
of the specimens, while France has about 5 percent and 10 per-
cent, respectively.) The 21 countries that lead in total number
of staff are among the 22 countries that lead in total number of
herbaria, listed in Tables 3 and 4, although the ranking differs,
as can be seen in Table 4. Finland, included in the tables, has
30 curators and ranks 23rd in staff size, whereas Belgium, not
included in the tables, has 31 curators and ranks 22nd in staff

710 Proceedings of the Biological Society of Washington

size. Except for Belgium, therefore, Tables 3 and 4 include all
countries with 30 or more staff members as of 1963.

Few if any of the world’s herbaria would claim to be staffed
adequately, and almost every curator would consider himself
overworked, Yet there are no absolute standards by which one
may judge the adequacy of professional (or technical and cleri-
cal) staffing. Instinctively, one can say that any herbarium
with less than one full-time curator is understaffed or that any
person holding down two curatorial positions, as about 2 per-
cent of the world’s curators apparently do, is overworked. Say-
ing this hardly sheds light on the general question. There are,
however, two useful ratios that measure objectively the relative
adequacy of staffing of a herbarium or country: (1) average
number of specimens per curator, and (2) average number of
curators per herbarium. Thus herbaria or countries can be com-
pared with each other or with the world as a whole by their
specimen:curator ratios. Likewise, countries can be compared
with each other or with continents or the world as a whole by
their curator: herbarium ratios. To be sure, these ratios may bear
little relationship to the level of activity in particular cases,
especially where a significant fraction of the curators identified
with an institution or country are not actually engaged in her-
barium research and curation; nevertheless, these ratios are the
only objective measures of staffing we have. Other factors
being equal, an above-average curator:herbarium ratio reflects
a favorable staffing situation, while an above-average speci-
men:curator ratio, i.e., more than the average number of speci-
mens per curator, reflects an unfavorable staffing situation.

Average curator:herbarium and specimen:curator ratios
are given in Table 2 for continents or regions and Table 4 for
the 22 countries with the most herbaria. As already mentioned,
there is an average of 4 curators/herbarium among the 794
herbaria reporting staff. South America, as a continent, leads the
world with an average of 5.5 curators/herbarium, followed by
Europe with 5.0/herbarium. North America trails with 3.0/
herbarium. Among the 22 top countries, the Netherlands leads
with 10.9 curators/herbarium, while the United States trails

with 3.0/herbarium. The favorable South American ratio ap-

pears to reflect aggressive herbarium growth on this continent

Natural history collection symposium vals

and also a liberal concept of reckoning staff (see below). The
relatively high European ratio seems to be a more authentic rep-
resentation of the true situation.

The average number of specimens/curator among the 794
herbaria is about 47,000. The herbaria of the British Isles lead
the world with an average of almost 95,000 specimens /curator,
while continetal European herbaria follow with 78,000/cu-
rator. The lowest ratio is to be found in the West Indies, where
each man curates an average of about 6,000 specimens. Ignor-
ing Madagascar, where there are only 3 herbaria, the second
lowest average for a large region, about 10,000 specimens /man,
is found in South America. By country, Switzerland leads with
about 170,000 specimens/man, followed by Czechoslovakia
(134,000/man) and the USSR (87,000/man). The high ratio
of specimens to curators in Europe and the British Isles reflects
the existence here of old, very large herbaria. In general, the
European countries rank above average both in curators to her-
baria and in specimens to curators.

In North America, the United States, with about 50,000 speci-
mens/man, ranks near the world average, while Canada, with
19,000/man, and Mexico and Central America together, with
about 10,000/man, rank well below the world average. On the
basis of curators/herbarium, the United States, Canada, and
Mexico and Central America all rank below the world average
at 3.0, 3.2, and 3.3, respectively. Among North American her-
baria, therefore, those of the USA are the least well staffed.

As a group, the world’s largest herbaria appear to be se-
riously understaffed. The 17 herbaria with 2 million or more
specimens (Table 13) have among them 38 percent of the
world’s 148 million specimens but only 8 percent of the world’s
3,158 curators. The 11 herbaria with the largest professional
staffs (20 or more members each) have 19 percent of the
world’s herbarium specimens but only 11 percent of the world’s
curators. If one computes ideal professional staff size for these
11 herbaria on the basis of the worldwide average of about
47,000 specimens/curator, the results, given in Table 16, are
very interesting. By this standard some herbaria prove, as
expected, to be grossly understaffed, but others, surprisingly,
seem to be even more grossly “overstaffed,” if indeed one may

712 Proceedings of the Biological Society of Washington

speak of any herbarium being overstaffed. Certainly the staff
figures of individual herbaria must be regarded with some
skepticism and be interpreted in the most cautious, relative
terms, because of the lack of uniformity among institutions in
reckoning who is a professional staff member. Thus, for ex-
ample, the herbaria at Montpellier, Sao Paulo, and Buenos Aires
report essentially all faculties of their respective botanical
institutes instead of just those persons who actually might
be considered tc belong to the professional curatorial staff.
Probably Kew ranks first in number of authentic herbarium
staff, which means that the largest staff in 1963 totalled
about 40 professional persons. Despite individual discrepan-
cies, it is noteworthy that the 11 herbaria as a group have only
about half (56 percent) of the professional staff that they
should have to meet average conditions.

It may seem unfair to measure the adequacy of staffing in
large and small herbaria by the same specimen:curator ratio,
because the small herbarium must have a relatively larger staff
for its size than the large herbarium. The maintenance of any
herbarium, regardless of its size, entails certain basic curatorial
tasks and functions, and minimum staff size, obviously, is one
person. The larger the herbarium, the more efficient it becomes
in terms of number of specimens that a curator can manage.
Tending to counteract this gain in efficiency, however, is the
greater workload of the large herbarium, which gains in service
responsibilities to the scientist and layman as it gains in size
and thereby general usefulness and visibility. It is problemat-
ical, therefore, whether the large herbarium should be mea-
sured by a different specimen:curator yardstick than the small
herbarium.

For purposes of discussion I have assumed until now that
the more than 3,000 persons listed in I.H. are all professional
curators, because there has been no other firm basis on which
to analyze professional manpower in the world’s herbaria. In
fact, as already indicated (“Source of Statistics”), this is not a
safe assumption. While the I.H. figures may give a roughly ac-
curate picture of the number and deployment of the world’s
herbarium-affiliated botanists, although even this can be dis-
puted, given the kind of peripheral scientific staff that one

Natural history collection symposium AS

finds listed for some herbaria, quite clearly the more than 3,000
persons who are listed for the 794 herbaria reporting staff do
not all engage actively in curatorial work or work that can be
construed as contributing directly to the building and mainte-
nance of these herbaria. Many are associated researchers or ad-
ministrators who have few if any routine curatorial responsibili-
ties. This is not to denigrate the essential, if sometimes indirect
or intangible, contribution of such personnel to the well-being
of the collections and the scientific life of the herbarium. Yet it
should be recognized that probably no more than 1,500 to 2,000
of the approximately 3,000 staff are really curators. The extrap-
olated figure for all 933 herbaria would be 1,800-2,400 cu-
rators.

Finally, professional staff represent only part of the world’s
herbarium manpower. To their number must be added the
technical and clerical supporting staff. At the U. S. National
Herbarium, the ratio of supporting to professional staff has
tended in recent years to remain at about 1:1. This certainly
is neither the best nor the worst ratio among the world’s her-
baria. If for comparative purposes we may assume that it is an
average ratio, then all figures given for professional staff should
be doubled to project total herbarium manpower. It seems likely
that upwards of 7,500 persons, working in one capacity or an-
other, are employed in the 933 herbaria treated in J.H. Consid-
ering that these 933 herbaria could represent a fifth or less of
the world’s public institutional herbaria (see “Source of Statis-
tics”), one must conclude that at the least there must be well
over 10,000 persons employed in herbarium-related work ( pro-
fession, technical, or clerical) and at the most there could be
upwards of 35,000 or even more persons manning the world’s
herbaria. Probably the truth lies somewhere between these ex-
tremes.

THE MOopERN PREDICAMENT
On 8 May 1964, a symposium was convened at Michigan
State University on the theme “The Herbarium in the Modem
University” to dedicate new quarters for the university's herbar-
ium, founded in 1863. (These quarters, in a renovated old build-
ing, had been occupied since the summer of 1963.) The event
was a resounding success. On short notice, 160 persons, rep-

714 Proceedings of the Biological Society of Washington

resenting 49 institutions, attended. Thus the taxonomic com-
munity responded to this herbarium pulse-taking with a vital-
ity that few would have predicted. Later, when the symposium
was published, John H. Beaman, curator of vascular plants and
organizer of the symposium, could write (Beaman, Rollins, and
Smith 1965, p. 113), “The attention which the program at-
tracted was an effective demonstration of the high level of cur-
rent interest in the herbarium as a resource for taxonomic
teaching, research, and service.” As local administrators said
convincingly what their subordinate curators wanted to hear
and as the speakers optimistically tallied up several hundred
years of achievements, pointing to unprecedented growth and
activity at present, those attending found themselves engulfed
in a euphoria of hope and prosperity. The National Science
Foundation, indispensable patron of American science, was
duly represented by the director of the Systematic Biology Pro-
gram, who then was Walter H. Hodge, himself a botanist. Dr.
Hodge presided and, while acknowledging such chronic and
worrisome problems as inadequate public understanding, fi-
nancing, staffing, and facilities, was able to conclude his sum-
mation on an upbeat with the welcome appraisal that the her-
barium today is “progressing rather than regressing.”

During the first week of September 1968, just five years after
the renovated building had been occupied, a demolition crane
moved into position, and its great iron ball began swinging.
In exactly one-half day, less time than it took for the dedication,
the building that was opened with fanfare and great hopes in
May 1964 was reduced to a pile of rubble! The pendulum of
progress had swung, pulverizing a modern university herbar-
ium “to make way,” in the words of Fortune magazine writer
Duncan Norton-Taylor (1967), “for the driveway to the new
Administration Building.” For the second time in six years the
whole collection of plants had to be moved, at last to truly new
quarters, but again at great cost in effort and lost research time.

To be sure, I have not told the whole truth. The curators
knew when they first occupied it that this newly renovated
building could serve only as an interim home for the herbarium
during the indefinite period between vacating the original
quarters and moving into some permanent quarters yet to be

Natural history collection symposium 715

planned and built. They did not know how very temporary the
interim quarters were to be. Now the herbarium is located in
the recently built Plant Biology Laboratories, where it occupies
twice as much floor space as it had occupied in its original
quarters. So well off is the herbarium, in fact, that for the first
time its fortunes are even cause for a certain amount of envy
at the university.

But is this momentary good fortune illusory? The present
quarters also are a temporary reftuge—hopefully for no more
than 10 years. A truly permanent home is to be provided some
day in a new museum building not yet begun. Furthermore,
the two curators who are mainly responsible for developing and
maintaining the herbarium (Beaman and H. A. Imshaug, cu-
rator of cryptogams)* must run a full research and teaching pro-
gram for graduate and undergraduate students while also trying
to manage a collection of more than 200,000 specimens, to
which are accessioned about 10,000 specimens/year. This is
100,000 specimens/man, twice the national average, and at this
rate of accessioning the herbarium should be gaining a new pro-
fessional staff member every 4-5 years.

The Michigan State University Herbarium certainly is
not impoverished; neither are the responsible university ad-
ministrators myopic. Quite to the contrary, it is a university
herbarium of unusual vitality with indefatigable curators and
with administrators who thus far have demonstrated uncom-
mon understanding and foresight. Yet this is precisely the
point: the university herbarium today (indeed the herbarium
in general) seems at best to lead a fragile existence, and no
amount of activity and leadership can cover up the ever-present
stresses and strains that threaten this existence constantly. As
Beaman (1965, p. 113) writes, “The herbarium is the oldest,
most essential, most expensive, and most difficult to develop of
all facilities for the study of systematic botany. Consequently,
the occupancy of new quarters by a herbarium, however mod-
est, is an event of note.” Wrapped up in his words is the para-
dox of the herbarium, especially in the university setting: es-

9 This is a good case in point of how the number of staff listed in Index Herbar-
iorum may bear little relationship to the number actually responsible for most or all

of the curating. Of the 10 persons listed, only 2 (Beaman and Imshaug) were, as
of 1963, carrying much of the burden of curation.

716 Proceedings of the Biological Society of Washington

sential but too expensive to be developed, accommodated, and
maintained adequately. The elements of crisis or collapse,
namely, collections that continually are outgrowing facilities,
staff, and other resources and a science that constantly is chang-
ing, are always present. The slightest erosion, therefore, of the
historic scientific and intellectual foundations of the herbar-
ium can precipitate instant crisis, and this is what we seem to
be witnessing with increasing frequency as classical botany
comes under the molecular gun. Confused by challenges of the
scientific worth of the herbarium, administrators may need lit-
tle persuasion to decide that the herbarium is an expensive, lat-
ter-day white elephant, which in terms of resources demanded
is a facility that drains more than it adds to a modern science
program.

The Michigan State University symposium dealt only with
university herbaria. In the international commerce of taxo-
nomic research, however, the large nonuniversity herbaria are
crucial institutions. What then is the state of affairs in such
large herbaria as the U. S. National Herbarium at the Smith-
sonian Institution? Today, with 3 million specimens, it is one of
the ten largest herbaria in the world and one of the three larg-
est in the New World. The bulk of these specimens has been
accumulated during the present century. As Stern (1966, p. 8)
has said, “it is a safe assumption .. . that there is no serious re-
search of any scope which can be executed in systematic botany
in the United States without some recourse to the plant speci-
mens of the U. S. National Herbarium.” One might amend this
statement by saying that any taxonomist in the world wishing
to conduct serious research on temperate North American
plants surely will need to take recourse to collections of the
U.S. National Herbarium, among others in the United States,
at some time during his study.

By some standards the U. S. National Herbarium has often
seemed the rich uncle among herbaria in the United States. As
the largest of the few American herbaria with direct access to
appropriated federal funds, it appears to occupy a favored posi-

19

tion.!” During the past few years the Smithsonian’s Depart-

10 Direct appropriation is not the only form of federal support in the USA. Since
the National Science Foundation was formed, large, though inadequate, amounts of

Natural history collection symposium Tf

ment of Botany has indeed experienced unprecedented growth
and prosperity. In 1965, the department and its herbarium
were able finally to occupy new quarters, a move culminating
years of dreams. A year later hopes were raised (Stern 1966 )
for the acquisition of new metal cases to replace the more than
2,200 archaic, inefficient wooden cases, which are not insect-
proof. For the first time in history the National Herbarium
seemed to be heading toward a fully modern facility, even if,
as at Michigan State University, the quarters were hardly de-
signed for a herbarium (e.g. a plant-drying facility was not in-
cluded in the plans! ). The staff of full-time professional bota-
nists had grown to an all-time high of 16.

Already this hard-won improved status has begun to erode, as
the inexorable growth of the collections continues without a
concomitant increase in space and staff. At present, in fact, the
department has a smaller full-time professional staff (13)*! and
less available office space (1.2 rooms/man instead of the orig-
inal 2/man), which is occupied to the point of crowding, than
when it moved in 1965. Although the specimens per curator
ratio is only one index of staffing adequacy, yet it is significant
that on this basis the department should have about five times
its present number of full-time curators, to say nothing of sup-
porting staff (Table 16), just to meet average conditions. The
professional botanists continue to do much of the routine cu-
ratorial work because of the perennially unfavorable ratio of
curatorial assistants to curators which temporarily may reach
as high as 1:2 but usually is 1:3-4. Owing largely to understaft-
ing, some 200,000 specimens, as many as the Michigan State
University Herbarium comprises altogether, must remain in
dead storage, freezing nearly a fourth of the available storage
cases. Specimen storage space probably will reach saturation
conditions in the herbarium in less than 10 years, by which time
federal support have been granted to many American herbaria for research and facili-

ties. This fact sometimes is overlooked, and the myth arises that the National Herbar-
ium is the only federally supported herbarium in the USA.

11 This number, unlike the figure of 21 in I.H., excludes resident emeritus curators,
honorary research associates, collaborators, postdoctoral associates, and long-term
visiting scholars who usually swell the professional ranks by 10—15 persons a year but
do not have obligatory curatorial responsibilities, although frequently they contribute
much help. It includes, however, several full-time staff botanists of the department
who have little or no responsibility for the collections.

718 Proceedings of the Biological Society of Washington

virtually all working space in the herbarium will be occupied
by cases. Some parts of the herbarium already are so over-
crowded that specimen filing is difficult if not impossible. The
effort to replace the wooden cases in toto collapsed, and as of
today only a relatively few have been replaced.

Meanwhile, the National Herbarium continues to be very ac-
tive, and the workload only increases. Over the 10-year period
from 1958 to 1967, about 700,000 incoming specimens—41,000
to 120,000/year and averaging 70,000/year—have been proc-
essed as gifts, exchanges, or specimens collected by or for the
herbarium’s botanists. During the same period, 20,000 dupli-
cate specimens/year of the 70,000 have been turned around and
sent out on exchange, while 37,000 specimens/year have been
mounted for addition to the herbarium, leaving about 13,000/
year that of necessity have gone into dead storage. Thus some
50,000 specimens have been retained, which means that every
year the herbarium should be adding at least one new profes-
sional botanist and commensurate supporting personnel just to
cope with the intlow and processing of material. Incoming ex-
change has averaged 25,000 specimens/year, leaving an ac-
cumulating exchange deficit of 5,000/year. Duplicate exchange
usually is a deficit operation for the large herbarium, which by
virtue of its size and importance must cooperate in many more
exchanges than the small or medium-sized herbarium. To at-
tempt to balance the books is futile: the more specimens sent
out, the more that come back, and the total inflow always seems
to outstrip the outflow. If, therefore, the National Herbarium
suddenly were able to find the extra 5,000 exchange duplicates
each year to meet the deficit, any balance would only be mo-
mentary, because the cooperating institutions would be stimu-
lated quickly to send us still more specimens, perhaps doubling
or tripling our annual deficit. Loans for research also have in-
creased steadily from the 16,700 specimens borrowed from the
National Herbarium in 1961 to the 41,500 sent out in 1967,
averaging about 25,000/year over the 1958-67 period. Finally,
requests for identifications keep rising, and nearly 180,000
identifications were made over these 10 years. In short, the
National Herbarium, like any large herbarium, is big business.

To a greater or lesser extent, nearly every herbarium in the

Natural history collection symposium 719

world is faced with the problems, dare I say predicament, of
the Michigan State University Herbarium or the U. S. National
Herbarium (e.g., see Rollins et al. 1967-68). Regardless of the
category of transaction, there seems to be no way to stem the
rising workload and service demand. At the same time the in-
tellectual foundations of the herbarium seem to be crumbling
within science today with an ever-increasing tempo making it
harder and harder for herbaria to justify and secure the kind of
support needed. Given the intensifying predicament, serious
crisis cannot be far away. It is regrettable, therefore, that in-
ner-circle conclaves like the 1964 symposium do not, for all
their timely challenges and encouragements, challenge any of
the sacred cows or age-old premises of the herbarium mentality.
The handwriting, it would seem, is on the wall, and the mes-
sage should cause concern if not alarm. If through rosy glasses
a move to new quarters means growth and prosperity, plain
sight might reveal that it really means harassment and retrench-
ment, with the collections being chased from one temporary
asylum to another, never gaining a permanent berth in their
own right and always being put out of mind administratively by
another wishtul promise. It is becoming critical, surely, for cu-
rators to interpret the signals correctly.

Clearly it is time to establish new relevancies and strategies
for the herbarium. Considering that herbarium growth poten-
tially is limitless, it is not surprising that the kind of statistics
cited above give administrators uneasy feelings. Unless there
are new objectives with rational limits and strategies that go
beyond merely asking for bigger and better facilities, the cur-
rent predicament is likely to deepen into an insoluble crisis, lo-
cally and generally.

Economics Or HERBARIA

Investments and Costs

To my knowledge, a thorough analysis of capital investment
and cost of operation has never been made for herbaria. This
important task will require lengthy study to produce complete
and reliable results, and herbaria, at least on a national basis,
should attempt it as a basis for seeking more federal support. It

720 Proceedings of the Biological Society of Washington

is, in fact, an almost impossible task, given the great variation
of facilities and expenditures from one herbarium to another,
not to mention the problem of currency differences between
countries. My cost analysis, which is rough and sketchy, is based
on extrapolation from the situations at the Michigan State Uni-
versity Herbarium and especially at the Smithsonian’s U.S. Na-
tional Herbarium. I may be presumptuous to attempt this, but
surely some hints of costs are needed.

The cost of herbarium space and equipment is ap-
proximately $50/sq. ft. at Michigan State University and
approximately $100/sq. ft. at the Smithsonian Institu-
tion, where, however, the density of stored specimens per
square foot is about double that of Michigan State. Con-
sequently, the static cost of housing specimens is about $2/speci-
men in both cases. Projected on a national scale at this rate, the
capital investment for herbaria in the United States is at least
$70 million today, and the worldwide investment is nearly a
third of a billion dollars. Even if the average cost were only
$1/specimen the worldwide investment would be $150 million.

In any herbarium, the specimen storage cases are the main
item of equipment. The U.S. National Herbarium housed its 3
million specimens as of 1963 in about 2,200 cases.'" Figured at
$100/case, which was the minimum cost of replacement at that
time, these 2,200 cases represented an investment of almost a
quarter of a million dollars. Using the National Herbarium’s
average of about 1,350 specimens/case, one can extrapolate,
and on this basis the USA had some 25,000 cases as of 1963,
while there were about 110,000 in the world. (Phanerogamic
specimens average only about 1,000/case, whereas some of the
cryptogamic groups average more than 1,350/case.) At $100/
case or its equivalent in other currencies, these totals represent
investments of about $2.5 million in the USA and about $11 mil-
lion in the world.

Overall operating expenditures vary from year to year and
herbarium to herbarium. One of the problems of cost estima-
tion, given the budget of a herbarium, is to separate research
costs from curatorial and herbarium-service costs. Thus, for ex-
ample, the Smithsonian’s Department of Botany operated with

'2 Fach case has 24 compartments,

Natural history collection symposium veal

about $400,000 in Fiscal Year 1968 (July 1967—June 1968), in-
cluding granted as well as appropriated funds, which averages
over $0.12/specimen for the approximately 3.25 million speci-
mens on hand by this time. These funds covered salaries and
operating funds for all research and curatorial activities, how-
ever, and probably no more than half of the total sum, i.e.,
about $0.06/specimen, was expended to support the U. S. Na-
tional Herbarium per se. For Fiscal Year 1968, therefore, one
might extrapolate that the USA spent at least $4 million on the
nation’s herbaria, including research and curation, and that at
least $2 million of this went directly to the support of herbarium
curation and service. The comparable figures for the whole
world would be nearly $20 million and $10 million, respectively.

The routine operation of a herbarium includes accessioning,
loaning and borrowing, exchanging, sorting and filing newly
mounted specimens, identifying plants, answering public en-
quiries, and other activities. Figures on a few of these opera-
tions will indicate how rapidly the expense of operating a her-
barium mounts.

The cost of sorting and filing newly mounted specimens var-
ies greatly, depending especially on the training and experience
of the person who does the work. Other things being equal, a
professionally trained botanist can sort and file much more
rapidly and efficiently than a technical assistant, but in either
case the speed and efficiency are direct functions of experience.
The botanist will earn two or three times more money per hour
and should, therefore, be three or four times more efficient than
the technical assistant, but this is not likely because the bota-
nist will file less mechanically and will take time out to solve
more problems. At the U.S. National Herbarium, where sort-
ing and filing are shared by botanists and assistants, it costs a
minimum of $0.10/specimen and an average closer to $0.15/
specimen for the whole process, which, for 50,000 specimens,
year, represents an annual bill of $5,000-$7,500 or even more.
Extrapolating on the basis of $0.15/specimen and assuming
that the annual growth rate of collections is about 1.5 percent
(see “Size of Collections”), one can estimate that the yearly
cost of sorting and filing newly mounted specimens is a mini-

mum of $75,000 in the USA and $330,000 in the world. These

722 Proceedings of the Biological Society of Washington

calculations also assume, for the sake of argument, that all
newly accessioned specimens are being mounted and filed
promptly.

Loan transactions constitute big business at the U. S. Na-
tional Herbarium, where today about 1 percent of the total
collection goes out on loan in a year. Personnel of different
levels are required to process these loans, but the total cost, by
my estimation, is equivalent to a professional man-year at
$13,000-$15,000. Thus the cost averages upwards of $0.50/
specimen. Extrapolating, the annual rate of loaning would be
about 350,000 specimens in the USA, costing about $175,000,
and about 1.5 million in the world, costing about $750,000.
These are, of course, very rough estimates.

The cost of public service is, like all other activities of the
herbarium, difficult to estimate. One important facet of public
service is plant identification. During the most recent 10-year
period for which there are statistics, the U. S$. National Herbar-
ium averaged about 18,000 identifications /year for professional
and lay persons. This represents less than half of the requests
actually made. At a very minimum this identification service
has cost $1/specimen, and a more realistic average figure would
be at least $2-$3/specimen. The rate depends on the percent-
age of the identifications made for professional persons, who re-
quire an authoritative precision not required by the public. By
the time he consults both the collections and the literature, a
botanist not infrequently spends an hour or two on a single
specimen; therefore, the cost can mount quickly to $5-$20/
specimen. In recent years, some Smithsonian botanists have
identified up to 4,000 specimens/year, mostly for professional
colleagues. Taking the rock-bottom figure of $1/specimen and
the rate of identification of the National Herbarium, I estimate
the annual bill for the USA to be something over $200,000 and
for the world about $1 million. The true costs are probably
double these figures at least.

Dividends
One would be foolish to attempt to put a dollar figure on the

full value of the herbarium to science and society, because in
a very real sense this value is incalculable. At the same time,

Natural history collection symposium 7123

herbaria do cost big money, as we have seen, and the public has
the right to ask, as it frequently does, what the payoff is. Cura-
tors are justified, therefore, if not duty bound, to consider what
dividends can be reaped from their collections and activities.

The worth of the herbarium to the scientific community can
be evaluated in part by the amount of money invested in her-
barium-based research. With respect to the United States, some
interesting data on research investment can be found in the
statistics of the National Science Foundation. Over the last six
years (1963-68), the NSF, through its Systematic Biology Pro-
gram, has awarded grants totalling $10,653,500 for studies in
systematic botany (excluding viruses and bacteria). This is an
average of almost $1.8 million/year, and during the last two
fiscal years (1967, 1968) the amount awarded has averaged
about $2 million/year. Of the money awarded, about 40 per-
cent ($4.25 million ) has gone to floristic and monographic stud-
ies, which are vitally dependent on the herbarium, while
another 40 percent has gone to studies that are much less de-
pendent on the herbarium but are likely to require it at some
stage just the same. In other words, about 80 percent of the
money awarded has gone into researches that are to some de-
gree herbarium-based. This represents about $1.4 million/year
or, if only floristic and monographic researches are considered,
about $0.7 million/year. The above figures are based only on
grants made by the Systematic Biology Program. It must be
added that environmental and other biologists who receive
grants through other NSF programs frequently conduct re-
searches that require the use of the herbarium. The money
spent through the Systematic Biology Program represents,
therefore, only the most direct and visible of NSF's investments
in herbarium-based research.

Much herbarium-based research is done in the United States
each year without financial support from the NSF. There prob-
ably are about 1,000 plant systematists in the United States. On
the basis of the past two years the NSF would seem to be sup-
porting only about 14 percent of these American taxonomists
(about 140 out of 1,000). (To the 80 new grantees each year
must be added about 60 continuing grantees; the average grant
lasts about 21 months. ) The 14 percent have been commanding

724 Proceedings of the Biological Society of Washington

nearly $2 million/year, but they constitute the “rich cousins” of
the taxonomic fraternity. Probably, the other 86 percent do not
average more than 10-15 percent of the almost $15,000/man/
year that the NSF-supported scientists have available for re-
search. At 10 percent or $1,500/man for the other 86 percent,
the annual investment for research in plant systematics in the
United States becomes $3.3 million ($2 million from NSF for 140
systematists + $1.3 million from other sources for 860 systema-
tists). The 10 percent estimate could be much too low, of course.
Carrying our extrapolation to its conclusion, we can estimate
that $0 percent of the $3.3 million, i.e., $2.64 million, goes into
herbarium-based researches of some type, while 40 percent, i.e.,
$1.32 million, goes to the support of floras and monographs,
which cannot be produced without the herbarium. Thus a $70
million herbarium investment supports $2.64 million worth of
research each year, although much more research could be sup-
ported annually with the same investment.

There is another means of evaluating the worth of the her-
barium to science in America. Given that the existence of the
herbarium is vital to the existence of the discipline of plant sys-
tematics, we can say that in the United States today’s 1,000 sys-
tematists are supported as a research fraternity by the $70 mil-
lion herbarium investment. At an average salary of $12,000/
year, the annual price tag of this fraternity is $12 million. Add
to this the $3.3 million used te pay for their research, and we
have a scientific enterprise costing $15 million annually that
could not exist as we know it today without the historical in-
vestment in the herbarium.

The cost of a service is also a measure of the value of the ser-
vice rendered. Thus in the previous section (“Investments and
Costs”) the costs to herbaria of storing and lending specimens
for research and of identifying plants for scientists and the pub-
lic, which probably are the two most important services of the
herbarium, are discussed.

The public user community is essentially the citizenship at
large, and its dependence on the herbarium can only be evalu-
ated in terms of specific kinds of requests such as for plant iden-
tifications. At any large herbarium, identification, like speci-
men lending, is big business. The annual bill for identification

Natural history collection symposium 125

would be much higher if the manpower were available to meet
the real demand.

CHANGING ROLE Or HERBARIUM

Historically, herbaria were the personal collections of private
individuals who preserved plant specimens to document cere-
monial or medicinal uses or to satisfy cultural or scientific curi-
osity. The collector, if a serious scholar, was both scientist and
curator. He traded duplicate specimens with colleagues as a
means of diversifying his own herbarium and of notarizing his
own finds. A man of means (e.g., of royalty ) could hire a cura-
tor and commission collectors to obtain the necessary speci-
mens for duplicate exchange, but his herbarium remained a
personal property for his own amusement and his curator’s, if
not his own, study.

The emergence of botany as a science in the 17th and 18th
centuries invested dried plant collections with a new signif-
icance and thereby brought about the institutionalization of
the herbarium. Private collecting has never ceased, of course,
but today the herbarium is highly institutionalized. Not only is
the herbarium an essential scientific institution, but in the or-
ganizational sense it has become a public institution, governed
by museums, universities, botanical gardens, and other cor-
porate bodies. The modern herbarium, in addition to being a
place of research, is a large service bureau. As already indi-
cated several times, herbaria in the aggregate represent big
business, and running a major herbarium calls for businesslike
methods. The private little collections that once could be known
in their entirety and be managed “out-of-pocket” by their sole
curators have become so massive in many cases that no single
curator could hope to know their limits or to discharge all their
tasks. Large herbaria require the cooperation of several to
many curators.

The fact is that although more than four centuries have passed
since the first institutional herbarium was formed curatorial
mentality and practice are still characterized strongly by the
personal entrepreneurship of a private collector. The transition
from the personalized, private herbarium to the collective, pub-
lic herbarium, with its corporate research and service responsi-

726 Proceedings of the Biological Society of Washington

bilities extending over many generations of curators and citi-
zens, has been made imperfectly at best in most instances.
Curators, whether they have curated a small herbarium or
some part of a large herbarium, have always tended to shape
their collections according to their own scientific and manage-
ment concepts. Often too little thought has been given to the
implications of being part of a much larger system that must
survive the lives and whims of individuals. Small, one-man
herbaria may be able to withstand the consequences of genera-
tion after generation of subjective curation, but large herbaria
must have objective standards or in time they become a hodge-
podge of curatorial idiosyncrasies. Thus, a large herbarium, in-
stead of being curated by a uniform, generalized system, may
be curated as several autonomous or semi-autonomous fief-
doms. Sometimes each fiefdom has its own familial and generic
concepts or filing system.

Some subjectivity is essential, of course, because in the final
analysis the herbarium is not only a facility and a resource but
also an instrument of taxonomic research. If the instrument has
shaped the science, so has the science shaped the instrument.
The science of systematic botany itself is changing, however,
and becoming less descriptive. As it becomes less descriptive,
it tends to become less subjective; therefore, the herbarium
should become less a subjective instrument of research and
more an objective source of information. Such evolution in
function demands new ground rules for collection building and
management.

The life cycle of plant taxonomy, whether one thinks of the
historical development of the science or of the knowledge about
a particular flora or group of plants, has had at least four rec-
ognizable phases thus far (cf. Valentine and Love 1958): de-
scriptive (exploratory ), floristic-phytogeographic, systematic,
and biosystematic (including chemosystematic). A fifth, eco-
systematic phase is just beginning. These are relative states
of progress in the development of taxonomic knowledge, of
course, and as such describe not only chronological stages in
time but also phases of activity going on simultaneously within
the taxonomic community at any given period of time. Collec-
tion building has tended to reflect this changing cycle of taxo-

Natural history collection symposium 127

nomic approaches, i.e., the character of the collections being
accumulated has been influenced by the type of taxonomy
being done. Obviously, there is no perfect system for arranging
the herbarium so that it will serve these five phases—or any
other phases—of plant systematics equally well, nor can a cura-
tor change the physical arrangement of his herbarium to con-
form to the latest thinking every time some new research fad
comes along. The bewildering array of systems and partial sys-
tems in use among herbaria today, which often are long since
outgrown or overgrown and of which no two seem to be alike,
stand like ancient shipwrecks as mute testimony to the naviga-
tional errors of past curators who tried to keep pace with the
times by arranging part or all of their collection according to
current taxonomic concepts, only to have these concepts change
faster than they could rearrange the specimens consistently.

The herbarium first became a scientific institution when tax-
onomic botany, indeed all of biology, was almost entirely de-
scriptive, and the principles of organization and use established
then have largely dictated practice ever since. Through the
years botanical exploration has been a chief stimulus for herbar-
ium-founding, witness the geographical shift of focus of new
herbarium development from Europe to North America and
Asia and thence to Africa and South America in phase with the
general exploration and development of these regions of the
world. During the descriptive-exploratory stage the herbarium
takes shape as a repository of exemplars of the new forms of
plant life coming off the collector's conveyor belt from exotic
regions. The descriptive or alpha taxonomist deftly and expertly
sorts from this conveyor, sifting the known from the un-
known. The known are filed and the unknown are described
and published as quickly as possible. At this stage the para-
mount function of the herbarium is to provide, for purposes of
identification and diagnosis, easy and logical access to the ex-
emplars of already-described taxa, and the primary task of the
curator-taxonomist is to keep his incoming material described
up to date, which requires that he know his previous collections
intimately and have them neatly classified and filed away.

The curator who likes to assign a place to every speci-
men has little difficulty in doing so while the herbarium is

728 Proceedings of the Biological Society of Washington

high on diversity and low on variability in its representation of
the plants in nature. As soon as a second specimen of a known
taxon appears, however, the exemplar approach begins to break
down, and each succeeding specimen further erodes the homo-
geneity of the taxon and complicates the task of identification
and novelty-recognition. Therefore, specimens additional to
the types not only have less value intrinsically than the types
but also constitute in reality a nuisance factor because they ob-
fuscate the nice boundaries that could be drawn on the basis
of single exemplars. The pigeonhole mentality is difficult if
not impossible to outgrow. For purely practical reasons every
specimen must have a place to rest, and, regardless of the phase
of taxonomic development, identification, comparison, and
diagnosis tend to remain the primary functions of the herbarium
and therefore dictate its arrangement. By the same token, the
herbarium botanist faces the danger of becoming trapped with
these functions, never having a chance to indulge in the broader
aspects of systematics.

Once the majority of the novelties have been discovered, at-
tention turns to floristics and phytogeography. In this second
phase of taxonomy the curator-taxonomist monitors the con-
veyor of incoming material for new and interesting distribution
records, and geography becomes a major parameter by which
he tries to sort and arrange his specimens. Now the currency of
study is not the taxonomic novelty, but the geographic novelty,
with endemism, disjunction, and the ebb and flow of floristic or
phytogeographic elements being major themes of interest. Gen-
erally, the curator-taxonomist will be especially interested in
only one or a few regions; consequently, his subdivisions will
be precise in these cases and very coarse for the rest of the
world. In time, such gerrymandered systems become clumsy
and meaningless as natural geographic arrangements; they also
become loaded with political anachronisms as the boundaries
of countries change. It probably is fair to say that hardly any
herbarium in the world uses a fully modern geographic scheme
of which it wholly approves. The geographic mentality fos-
tered by this phase of taxonomic development can lead easily
to absurd extremes in herbarium-packing of specimens of the
same species for purposes of documenting local distribution.

Natural history collection symposium 729

The third or systematic phase raises taxonomy and the use of
the herbarium above the level of pure description to the philos-
ophy of relationships. The herbarium now becomes an active
instrument of the curator-taxonomist as he tries to arrange the
specimens according to how they should be classified, and, as
mentioned before, most modern herbaria reflect some earlier
system of classification. A large herbarium hardly is amenable
to further manipulation as new systems are proposed. Further-
more, seldom is it possible even to keep current the older sys-
tem in use, if it is a phylogenetic one.

In the fourth, biosystematic phase, we see the need for large,
in-depth collections (population samples) of the taxa under
study. Most curators are justifiably reluctant to store large
samples of individual taxa because they cannot cope physically
with the specimens. Moreover, in terms of the traditional and
still prime functions of the herbarium, this represents uncon-
scionable duplication. Yet no biosystematist wants to see his
samples treated as “duplicates” and split up for inter-institu-
tional exchange. Also, the biosystematist needs a herbarium
that provides easy access to other kinds of specimen data than
the traditional name and place of collection.

The recent, ecological phase of systematics has only begun.
The next decade will bring, I believe, a solid alliance between
taxonomists and ecologists and the emergence of what can be
called “ecosystem taxonomy.” Surely the International Biolog-
ical Program (IBP) will develop intense pressures for this. The
ecosystem taxonomist, as contrasted with his predecessors,
will be less concerned with the absolute precision of his identi-
fications and the phylogenetic hierarchy of his organisms and
more concerned with the general, statistical patterns of distri-
bution as they correlate with environmental factors, including
pollutants; he will also be concerned especially with the inter-
relationships and coevolution of different plants and of plants
and animals, including man. Thus he will need a much more
flexible access to the data locked up in the herbarium than we
now have; furthermore, he will call for more sophisticated
ecological data-keeping.

The picture is clear. The herbarium was designed for the
purposes of a descriptive science that dealt mainly with the

730 Proceedings of the Biological Society of Washington

questions of what and where, but it has had to survive funda-
mental changes in this science and now finds itself in an era
when the questions are mainly how and why. The fact is that
the herbarium has never really adapted to the modern biosys-
tematic and ecological era, and unless it does it will become
largely irrelevant in time. We have not yet overcome the prob-
lem of providing flexible, multi-access to a data bank that can
have only one physical structure, ordered by one parameter, in
this case the scientific name. The sharp decline in the founding
of new herbaria since the 1920s seems only to be a specialized
reflection of a general decline in descriptive biology. The her-
barium is after all the chief resource of the descriptive plant sys-
tematist, and any deterioration of his status inevitably will de-
crease the demand for the tools of his trade. Collectors have
pushed to the limits of the temperate regions and pressed on
into the tropics. Perhaps, especially with new temperate and
tropical flora projects in progress and with greatly expanded
tropical exploration and research, the 1960s and 1970s will
prove in retrospect to have reversed the downward trend in de-
scriptive systematics and herbarium-founding, but this seems
doubtful, given the present-day climate of biology and science.
Despite the secondary resurgence of such activity particularly
in tropical regions, the downward trend appears to be inevi-
table and irreversible. The golden age of herbarium-found-
ing, has passed.

STRATEGY For THE FUTURE

As a physical creature, the herbarium has grown through
more than 400 years until today it has achieved menacing
proportions. Not just a few curators are virtually enslaved by
the sheer burden of the routine daily transactions and public
service, when in fact they should be practicing science. At the
same time the science, too, has changed, so that altogether the
forces of change and growth have conspired to make it diffi-
cult for today’s herbarium botanist to be both curator and scien-
tist. Descriptive taxonomy is a fairly natural and easy byprod-
uct of curatorial activities, and it thrives on a constant inflow
of new material. To the biosystematic, ecosytematic, or ex-
perimental taxonomist, however, curation is largely an encum-

Natural history collection symposium Tol

brance, a service to perform as the price of being a professional
taxonomist.

The herbarium, no less than the library, continues to fulfill an
absolutely vital role in science and in practical human affairs as
a data bank and information system, even though increasingly
it creaks from an overburdened, arthritic curatorial machin-
ery and suffocates in the clutch of the time-honored but out-
moded and inflexible ground rules of research and public ser-
vice. Being an institutional giant and in many respects an over-
aged one, it faces hazards of survival that are not small. There
are those today—and their number is growing—who see the
herbarium as an economic millstone and an intellectual dino-
saur in the modern scheme of science. The truth, however, is
that the herbarium is beginning to be tapped for a whole new
generation of scientific and public questions. As the concern
rises about the quality of our natural environment and the
ecological principles that control this quality, public officials
are being forced to come up with instant ecological histories
and forecasts. The conservation of natural resources, including
plant and animal communities and particularly endangered
species, has become a burning public concern. Ecology and
conservation quickly reduce themselves to relationships among
organisms. Museums, herbaria included, are the repositories of
vast amounts of raw and standardized data about the earth’s
organisms. The alert curator is not surprised, therefore, that
the rising emphasis on environmental biology is giving new sig-
nificance and urgency to the business of museums. Unfortu-
nately, herbaria, like museums in general, are not ready for the
increased demands of the era of environmental biology.

The time for new premises and strategies is upon us. The
principal challenge is to “get with it” in trying to reshape the
herbarium for the age of environmental biology and the com-
puter, to meet the contingencies not only of a changing science
( biosystematics, chemosystematics, ecosystem taxonomy, etc. )
but also of a moody, ecologically conscious society who want to
know how to survive. We must hope that the world’s herbaria
will unite at different levels (local, regional, national, interna-
tional) to develop a blueprint for action. Meanwhile, several
of the necessary steps to be taken are obvious.

732 Proceedings of the Biological Society of Washington

(1) Every herbarium is both a scientific organization and a
public service bureau, and the time has come to accept the
full import of this dual nature and reorganize accordingly. The
day is past when the taxonomic scholar can be both scientist
and curator. Our goal must be to isolate the functions of the
herbarium, which are the tasks of the curator, from the re-
search of the herbarium, which is the responsibility of the
scientist, i.e., taxonomic scholar. Only in this way can the her-
barium rise to meet the increasing service demands and at the
same time remain a viable scientific research institution.

The scientific and the service functions of the herbarium can
and should be performed by different staffs. As a public ser-
vice bureau, the herbarium should be organized like a modern
library and staffed by a cadre of professionally trained, librar-
ian-like technical experts and aides who specialize in the her-
barium’s functions, e.g., accessioning, filing, lending, identify-
ing, etc. Libraries are not organized on the premise that only
scholars can order, purchase, catalog, shelve, and loan the
books, and neither should herbaria be organized on this prem-
ise. After an overall systems and cost analysis of input, process-
ing, storage, and output, herbaria should departmentalize and
staff appropriately. Non-research personnel, whose professional
rewards do not depend on publication, can be trained to per-
form most if not all curatorial and public service functions of
the herbarium just as well as, if not better than, research scien-
tists. As a scientific organization the herbarium should become
an institute for advanced studies, organized and staffed accord-
ing to disciplines and programs, not by curatorial responsibili-
ties. A strong link and intimate cooperation should be main-
tained between the curators and the scientists, however, be-
cause the latter will need to continue to guide curatorial policy.

(2) The computer must be brought into the herbarium
without further delay. A new day has dawned in information
science, and the meaning of this for museums has been pointed
out repeatedly in recent years (Sokal and Sneath 1966, Squires
1966, Crovello 1967, Rogers et al. 1967, Soper and Perring 1967).
The constant growth of collections impels us to find more effi-
cient means of storing the specimens and accessing the data. A
computer system for information retrieval (IR) provides the

Natural history collection symposium 733

ideal answer to the problem of data access and allows great free-
dom in the physical arrangement of the specimens. Data can be
retrieved without necessarily taking recourse to the specimens,
and the cross-indexing power of the computer enables one to
find specimens when necessary regardless of the physical stor-
age system. The latter capability makes the computer an im-
portant tool for managing herbarium transactions (loans, ex-
changes, accessions, etc.) as well as for providing flexible
access to the embedded data. Various control lists can be gen-
erated that profile the strengths and weaknesses of the herbar-
ium with respect, for example, to geographic or taxonomic
representativeness of the collections. Such profiles could put
curatorial decision-making on a much more objective basis,
especially as regards the accessioning of new material.

Every specimen carries both objective data (e.g., geographic
and other label data), which may require little or no professional
interpretation, and subjective data (e.g., morphological traits ),
which may require highly professional interpretation that can
only be made after study of the specimen itself. In a manual
system, neither kind of data can be retrieved without actually
seeing the specimen, and this requires transporting either the
specimens to the investigator or the investigator to the speci-
mens. An IR system can bring the objective data from the
specimens to the investigator without burdening anyone with
handling the specimens themselves. Once a magnetic record
of a collection is created, one can in effect rearrange an entire
herbarium just to answer a single question and do it, perhaps,
with less effort and cost than to process a loan of a few hun-
dred specimens. Even with only a few descriptors per specimen
recorded, many combinations are possible, and one is able to
ask complex questions and thereby to locate precise subsets of
specimens or compile specific data from randomly scattered
places in the herbarium—all without moving a single specimen.
The investigator is free to decide on the basis of his answer
whether he needs to see the specimens. To be sure, there are
certain risks to retrieving and using data without seeing the
specimens; for example, the risk of misidentification. Neverthe-
less, there are many instances in scientific research and public

service when these risks are tolerable.

734 Proceedings of the Biological Society of Washington

Herbarium curators have in large measure lost control of
their vast data bank, now comprising an unmanageable 200
million specimens or more over the world. Given the advanced
state of computer technology today, there scarcely is a defense
any longer for continuing to add to this overburden of speci-
mens without simultaneously capturing the data for manage-
ment and retrieval. The high cost of developing and implement-
ing an electronic data processing (EDP) system will prohibit
indiscriminate input and force curators to make some hard deci-
sions about the specimens and data to be preserved. Thus the
process of computerizing data can serve as a much-needed qual-
ity control mechanism. If a specimen does not carry data worth
computerizing, then it can hardly be worth preserving and fil-
ing in the herbarium for all time. No longer can we afford to
presume on our successors by adding to their future curatorial
burden under the blithe assumption that while the specimen
was not worth our time and money it might be worth theirs.

The place to begin is with select subsets of our herbarium
collections (e.g., types) and with newly accessioned material.
It is doubtful whether herbaria will ever have the resources
to input the whole 200-million-specimen backlog, and, consid-
ering the quality of the data of many older specimens, one can
raise serious questions as to whether this should be done even
if the resources were available. Instead, we must concentrate
on what I might call the “forelog.”

(3) All herbarium operations need to be examined carefully
and modernized if necessary, not only with respect to EDP, but
also in the light of the current state of science, the growing
shortage of staff and space, and the growing public and profes-
sional demand for herbarium-based information. We have
seen that herbarium operations today are expensive; even a 1
percent increase in efficiency would effect significant savings.
Courage will be needed to abridge or abandon outmoded prac-
tices. To cite one prominent example, the time-worn specimen
exchange procedures badly need scrutiny and appropriate
streamlining. Particularly the larger herbaria need to shake
loose from the iron grip of the book-balancing exchange mental-
ity.

In the present day, duplicate exchange is in some respects an

Natural history collection symposium 735

anachronism. The world hardly lacks for large, representative
herbaria, and the number of herbaria has increased to the point
where it is difficult if not impossible for an institution to draw
rational limits to its exchanges. Today’s rapid means of travel
and communication leave little excuse for building up numer-
ous herbaria with duplicates of the same collections. In the
framework of population biology, moreover, it is questionable
whether there is such a thing as a “duplicate” specimen.

One way to modernize exchange practice would be to estab-
lish a cooperative exchange center or clearinghouse which man-
ages transactions by computer and provides specimen-sorting
service as needed. Each herbarium would trade and balance
books with the system, not with every herbarium from which it
happened to receive duplicates. The exchange center could
abide by the wishes of donors in distributing their sets of speci-
mens by sending them only to specified recipients. On the con-
trary, the center could honor the wishes of recipients by send-
ing them only the kinds of material they requested. In many
cases, specimens might be exchanged directly between donor
and recipient after clearing the transaction with the exchange
center. Such a system surely would reduce the inefficiency and
increase the effectiveness and order of the exchange process
and thereby serve the general good. It could only be established
by cooperative effort, however.

(4) The herbarium needs strengthened intellectual founda-
tions, including ties with some of the newer biological and en-
vironmental disciplines. This can be achieved by exploiting
the herbarium through EDP systems as a vital data bank for
biologists other than systematists and by organizing broad, pret-
erably interdisciplinary research programs that require herbar-
ium resources and data and at the same time engender the kind
of national and international planning and cooperation that
will give the world’s herbaria as a group more cohesion, sin-
gular and self-respecting voice, and 20th-century relevance.
Given the present experimental and molecular scientific cli-
mate, we can say that the herbarium is truly at the crossroads,
and it will surely fall if it loses its intellectual underpinnings
and fails to adapt to the communications revolution by com-

736 Proceedings of the Biological Society of Washington

puterizing. Literally, therefore, “united we stand and divided
we fall.”

(5) Finally, the herbarium needs a new, more solid finan-
cial base. This means greater national support, which can come
only through cooperative planning. Herbaria are a special kind
of national research archive and instructional tool, deserving of
regular, dependable subsidy. National support cannot be
sought on any rational basis, however, until herbaria show a
willingness to operate less like competitive, private enterprises
and more like the units of a functioning system. Every herbar-
ium that ever makes a loan or accepts an exchange or gift of
specimens is part of a national and international network, partic-
ipating in the commerce of taxonomy. Any government would
be less than prudent to sink large sums of money—and the
herbarium is a colossus that could absorb unlimited funds—into
its herbaria without first requiring conscious cooperation that
takes cognizance at a planning level of the actual network
formed by the existing herbaria and divides the responsibilities
so that duplication of resources and services is minimized.

In the United States federal support for herbaria over the
years has been trivial compared to the needs, and unfocussed
with respect to the national interest. The time is ripe for a na-
tional strategy. During the 6-year period from 1963 to 1968, the
Systematic Biology Program of the National Science Founda-
tion granted less than 2 percent of its funds for collection main-
tenance per se, and for botany this amounted to about $30,000/
year, totalling less than $200,000 out of $10 million for the
whole period. If a matching dollar had been put up for every
dollar spent on herbarium research, the amount would have
totalled at least $4 million and perhaps as much as $8 million—
or $0.7-$1.3 million/year. This would have brought an average
of about $15,000-$30,000 to every herbarium in the country
during 1963-1968. Surely, for every dollar spent on fieldwork
there should be a matching dollar for collection curation. Too
often research proposals in systematic botany take the availabil-
ity of herbarium collections and services for granted, when in
fact they should be asking for the funds to purchase this acces-
sibility and service just as they ask, for example, for funds to
pay publication costs. Ironically, curators as curators are prob-

Natural history collection symposium 137

ably the most thanked people in biology, but as individuals they
are taken for granted for the many thankless tasks they must
perform personally.

It would be unfair to imply that NSF support for herbaria
has been confined to the relatively few dollars that have come
directly through the Systematic Biology Program. In addition
there have been grants for facilities and more recently the Of-
fice of Science Information Services at NSF has been support-
ing the development of computer systems for biological data.
Nevertheless, the total investment has certainly not met the
needs.

Many American herbaria, especially in the smaller colleges
and universities, serve primarily a teaching function. Teaching
herbaria are vital to the nation’s science programs and should
be supported as educational facilities. Their subsidy should be
commensurate to the teaching programs they support. The
small teaching collection has a way of escalating into a larger
and larger research collection, demanding space and resources
that exceed the teaching value of the herbarium to its home in-
stitution. Such escalation may show commendable industry on
the part of the local curator, but it may also pose hard questions
about duplication of resources for the national funding agency
that is asked to pick up the tab.

At present the United States has only one National Herbar-
ium, but many other American herbaria receive federal support.
What is needed as a framework for greater federal support is
for all university, government, and private research herbaria in
this country to organize themselves into a full network of na-
tional herbaria. Then each could be recognized and financed
to fulfill a particular role. Some herbaria might concentrate on
broad geographic and systematic coverage, while others con-
centrate on particular regions or systematic groups. State and
local governments might be induced to support a complemen-
tary network of state and local herbaria, perhaps with some
matching federal funds if national scientific standards and ob-
jectives were being met. Any national system must steer a mid-
dle course between centralization and decentralization of re-
sources and management. The lesson of the Berlin herbarium,
destroyed in World War II, has taught taxonomists the wisdom

738 Proceedings of the Biological Society of Washington

of a certain amount of decentralization and duplication of im-
portant collections.

Part of the price of increased federal support, it is clear, will
be the loss of a certain amount of autonomy, because herbaria
will have to agree on their roles and stick to them. While there
may be no limit to the number of small teaching herbaria that
can serve a useful function, a nation only needs so many large
research herbaria of a given geographical or systematic spe-
cialty. The latter statement is especially true in a day when the
costs of transporting scientists or specimens may be cheaper
than the costs of maintaining essentially duplicate research her-
baria. Thus the further development of teaching and research
herbaria cannot be left to chance if the United States is to have
a national strategy for support. The whole point of a national
plan would be, not to compel anyone to do anything, but to
identify the role that each herbarium should play and then sub-
sidize the herbarium accordingly.

Procress Towarbd GOALS

The ideas presented here certainly are not new (e.g., see
Sokal and Sneath 1966). On an individual basis, many herbaria
have taken or attempted some of the steps proposed, in some
cases long ago. But the destiny of the herbarium as an institu-
tion can only be decided by cooperative planning and action.
Herbaria must unite to persuade national policy planners to
support them, and they must unite to train a pool of technical
experts who can take the burden of curation off the backs of
the scientists. Operational procedures like duplicate exchange
‘annot be modernized unilaterally. A herbarium only hurts
itself if it tries this. Likewise, no single herbarium has the
necessary resources or force of authority to develop and im-
plement a herbarium-wide data processing system. On the
international level, the International Bureau for Plant Taxon-
omy and Nomenclature, with headquarters in Utrecht, Nether-
lands, has done much to increase cooperation. Many of the
problems can be solved only by active collaboration at the na-
tional level, however. Eventually, such national collaboration
can lead to more formal international planning and cooperation.
Fortunately, at present a number of museum and herbarium di-

Natural history collection symposium 739

rectors in the United States have become quite concerned about
the future of museum collections and of systematic biology in
general, and two or three studies are underway to determine
the feasibility of greater national cooperation and financial sup-
port in the USA.

Within the last few years several pilot projects in applying
electronic data processing to museum collections have been
undertaken, which must be nurtured and expanded to embrace
all herbaria. The Smithsonian Institution now has programs
going to computerize data from selected portions of its col-
lection of about 60 million specimens. One of the projects,
headed by Mason E. Hale, is designed to produce an auto-
mated register of type holdings in the U. S. National Her-
barium. The cooperation of other herbaria is being sought
so that the Type Register can in time become a union list of
worldwide holdings. Another highly significant development is
the TAXIR (Taxonomic Information Retrieval) system and re-
search program of David J. Rogers and group at the University
of Colorado, financed by the National Science Foundation. Two
other noteworthy data-automation projects are under way at the
herbarium of the National Museum of Canada in Ottawa
(James H. Soper) and at the Herbario Nacional del Instituto
de Biologia, Universidad Nacional de México (Arturo Gomez-
Pompa).

North American plant taxonomists have just embarked on an
immense new cooperative program, Flora North America,
which holds much promise for bringing about greater herbar-
ium coordination. Flora North America is organized to pro-
duce a concise treatise of the vascular plants of the continent
north of Mexico, but the project is deeply committed to bring-
ing flora-preparation into the computer age. Information sys-
tems concepts are being exploited to develop a flora data bank,
which inevitably must involve the herbarium and the botanical
literature (Morse et al. 1968, Shetler 1969). Such a data bank
should help to translate the herbarium to many people and to
make it more relevant than ever as we enter the era of environ-
mental biology.

740. Proceedings of the Biological Society of Washington

CONCLUSION

The quantitative growth of the world’s herbaria has over-
whelmed us and become an end in itself, such that we spend all
of our time packing away specimens for a research day that
never comes. At the same time we find ourselves incapable
of retrieving the most elemental information. The time is here
if not past when a qualitative innovation in herbarium build-
ing and management is needed. To face the future the world’s
herbaria need a new strategy based squarely on electronic data
processing systems and a businesslike understanding of the ser-
vice and storage demands of the modern herbarium. The com-
puter makes possible a whole new concept of data banking in
the herbarium—the first real innovation since the specimen
case replaced the herbalist’s scrapbook—and the environmental]
biologist and ecosystem ecologists have already created the de-
mand for this kind of data access. The herbarium community
must unite in phrasing its needs and organize to meet and sup-
port them cooperatively. Today herbarium botany, like big
science generally, requires big money, and this calls for big but
responsible organizing premises and programs. Otherwise, the
herbarium ceases to be relevant in terms that the taxpayer or
even another scientist can understand.

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PA

Ciece, H. 1968. 4-stage MSU building started in 64, cost $4,050,000;
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Natural history collection symposium 741

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Lanjouw, J., AND F. A. StarLeu (compilers). 1954. Index herbariorum,
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Lawrence, G. H. M. 1951. Taxonomy of vascular plants. Macmillan
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NATIONAL SCIENCE FounDATION. 1967. Grants and awards for the Fiscal

742 Proceedings of the Biological Society of Washington

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SHETLER, S. G. 1963. Botany—A passing phase? AIBS Bulletin 13: 23-
20:

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———— In press. An information storage and retrieval system for bio-
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VALENTINE, D. H., Anp A. LOve. 1958. Taxonomic and biosystematic

categories. Brittonia 10: 153-166.

743

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748 Proceedings of the Biological Society of Washington

TABLE 5. Oldest and Youngest Herbaria.

Year of Number of
Location Founding Specimens
I. Founded before 1750
1. Padua (PAD), Italy 1545 229,000
2. Rome (RO), Italy 1566 400,000
3. Bologna (BOLO), Italy 1567 —
4. Leiden (L), Netherlands 1575 1,800,000
5. Basel (BAS), Switzerland 1588 200,000
6. Paris (P), France 1635 5,000,000
7. Paris (PC), France 1635 1,200,000
8. Floriana (ARG), Malta 1675 10,000
9. Siena (SIAC), Italy 1691 492
10. Amsterdam (AMD), Netherlands 1700 175,000
11. Torino (TO), Italy 1729 50,000
12. Reduit (MAU), Mauritius SY, 13,500
13. Vienna (Wien: W), Austria 1748 2,500,000
Il. Founded in 1959-631

1. Addis Ababa (ETH), Ethiopia 1959 6,000
2. Tampa (USF), Florida, USA 1959 48,000
3. Victoria (SCA), Cameroun 1959 1,400
4. Arcadia (LASCA), California, USA 1960 25,000
5. Heidelberg (HEID), Germany 1960 —_—_—_—_
6. Monrovia (LIB), Liberia 1960 2,500
7. Trieste (TSB), Italy 1960 11,000
8. Baghdad (BUA), Iraq 1962 5,000
9. Nsukka (UNN), Nigeria 1962 —_—__—
10. Port Victoria (SEY), Seychelles 1962 500
11. Aarhus (AAU), Denmark 1963 25,000
12. Brasilia (UB), Brazil 1963 20,000
13. San Marino (HNT), California, USA 1963 500

1 All of these but the Tampa herbarium first appeared in Index Herbariorum with
the 5th edition (1964); the Tampa herbarium was reported in the 4th edition (1959)
but without data.

Natural history collection symposium

749

TaBLeE 6. Herbaria Founded in Europe since 1950 (1951-63).1

Year of Number of

Location Founding Specimens

1. Berlin (BSP), Germany 1955 3,200
2. Avon (ABT), France 1958 20,000
3. Funchal (MADJ), Madeira 1958 1,500
4. Heidelberg (HEID), Germany 1960 ——
5. Trieste (TSB), Italy 1960 11,000
6. Aarhus (AAU), Denmark 1963 25,000

1 The earliest 12 herbaria to be founded in Europe are listed in Table 5.

TABLE 7. Oldest and Youngest Herbaria of the British Isles.

Re

Year of Number of
Location Founding Specimens
I. Founded before 1850

1. London (BM), England 1753 oo
2. Cambridge (CGE), England 1761 450,000
3. Edinburgh (E), Scotland 1761 2.,000,000
4. Glasgow (GL), Scotland 1780 170,000
5. Dublin (DUB), Ireland 1790 25,500
6. Bristol (BRISTM), England 1820 13,400
7. Manchester (MANCH), England 1821 3,000,000
8. York (YRK), England 1822 8,000
9. Norwich (NWH), England 1825 20,000
0. Warwick (WAR), England 1836 6,000
1. Torquay (TOR), England 1844 ————
2. Ipswich (IPS), England 1846 15,000

II. Founded since 1950 (1951-63)
1. Exeter (EXR), England 1953 25,000
1,000

2. Keele (KLE), England 1955

750 Proceedings of the Biological Society of Washington

TaBLe 8. Oldest and Youngest Herbaria of North America and the
West Indies.!

Year of Number of
Location Founding Specimens
I. Founded before 1850
1. Winston-Salem (SC), North Carolina lee 600
2. Charleston (CHARL), South Carolina 1773 ——_——_—
3. Middlebury (MID), Vermont 1800 3,000
4. Philadelphia (PH), Pennsylvania 1812 1,000,000
5. Montreal (MTMG), Quebec 1820 55,000
6. Philadelphia (PHIL), Pennsylvania 1821 —_—__—
7. Geneva (DH), New York 1822 16,000
8. Boston (MCP), Massachusetts 1823 14,000
9. West Chester (DWC), Pennsylvania 1826 12,000
10. Amherst (AC), Massachusetts 1829 84,000
11. Albany (NYS), New York 1836: 430,000
12. Ann Arbor (MICH), Michigan 1838 1,000,000
13. Saint John (NBM), New Brunswick 1842, 15,000
14. Madison (WIS), Wisconsin 1849 360,000
II. Founded since 1950 (1951-63)

1. Calgary (UAC), Alberta 1951 12,000
2. Sioux Falls (AUG), South Dakota 1951 1,000
3. Calgary (CFB), Alberta 1952, 5,000
4. Halifax (NSPM), Nova Scotia 1952 oo
5. Ottawa (CCO), Ontario 1952 10,000
6. Quebec (QFB), Quebec 1952 12,000
7. Baton Rouge (LSUM), Louisiana 1954 5,000
8. Saint John’s (NFLD), Newfoundland 1954 8,000
9. Chicago (CHI), Illinois 1955 20,000
10. Swarthmore (SWC), Pennsylvania 1955 4,200
11. Mayaguez (FPDB), Puerto Rico 1958 4,000
12. Saskatoon (SAFB), Saskatchewan 1958 2,000
13. Tampa (USF), Florida 1959 48,000
14. Arcadia (LASCA), California 1960 25,000
15. San Marino (HNT), California 1963 500

1 No herbaria were founded in Mexico or Central America during these years.

Natural history collection symposium 751

TABLE 9. Oldest and Youngest Herbaria of Asia.

Year of Number of
Location Founding Specimens
I. Founded before 1900
1. Calcutta (CAL), India 1793 1,000,000
2. Dehra Dun (DD), India 1816 300,000
3. Peradeniya (PDA), Ceylon 1817 85,000
4. Eskisehir (ESK), Turkey 1832 725
5. Coimbatore (MH), India 1874 124,525
6 Tokyo (TH), Japan 1875 10,000
7. Sapporo (SAP), Japan 1876 130,000
8. Sapporo (SAPA), Japan 1876 85,000
9. Tokyo (TI), Japan 1877 500,000
10. Hong Kong (Hk), Hong Kong 1878 30,000
11. Simonoseki (YAM), Japan 1883 12,000
12. Tomsk (TK), USSR 1885 357,000
13. Rawalpindi (RAW), Pakistan 1893 60,000
II. Founded since 1950 (1951-63 )

1. Istanbul (ISTO), Turkey 1952 2,800
2. Allahabad (BSA), India 1955 7,072
3. Yokohama (YNU), Japan 1955 10,000
4, Dehra Dun (BSD), India 1956 29125
5. Poona (BSI), India 1956 85,000
6. Shillong (ASSAM), India 1956 70,130
7. Tokyo (MAK), Japan 1958 380,000
8. Baghdad (BUA), Iraq 1962 5,000

752 Proceedings of the Biological Society of Washington

TABLE 10. Oldest and Youngest Herbaria of Australasia and the Pacific
Islands.

Year ot Number of
Location Founding Specimens
I. Founded before 1900
1. Bogor (BO), Indonesia 1817 1,000,000
2. Melbourne (MEL), Australia 1857 1,500,000
3. Wellington (WELT), New Zealand 1865 200,000
4. Christchurch (CANTY), New Zealand 1867 20,000
5. Singapore (SING), Malaya 1875 400,000
6. Adelaide (AD-U), Australia 1875 Sse
7. Brisbane (BRI), Australia 1880 500,000
8. Auckland (AKU), New Zealand 1883 12,500
9. Honolulu (BISH), Hawaii 1889 160,000
10. Rydalmere (DAR), Australia 1890 9,000
11. Kuching (SAR), Sarawak, Borneo 1895 25,000
12. Sidney (NSW), Australia 1896 750,000
Il. Founded since 1950 (1951-63 )
1. Adelaide (AD), Australia 1954 150,000
2. Alice Springs (NT), Australia 1954 15,000
3.

Laguna (CLP), Philippines 1954 1,000

53)

b

—_

Natural history collection symposium 753

TaBLeE 11. Oldest and Youngest Herbaria of Africa.

CUR © bo

a :
em SPSS ee plier) Sie Se Ne) i

Year of Number of
Location Founding Specimens
I. Founded before 1900
Cape Town (SAM), South Africa 1855 —_—_——_
Cape Town (BOL), South Africa 1867 137,000
Durban (NH), South Africa 1882 89,000
Grahamstown (GRA), South Africa 1889 100,000
Kampala (KAW), Uganda 1898 25,000
II. Founded since 1950 (1951-63 )

Cairo (CAIH), Egypt 1951 16,000
Lourenco Marques (LM), Mozambique 1951 20,000
Mahalapye (MAH), Bechuanaland 1951 1,345
Salisbury (CAH), Southern Rhodesia 1955 12,000
Dedza (NYAS), Nyasaland 1956 4,500
Elisabethville (EBV), Congo 1956 30,000
Rabat (RAU), Morocco 1957 ————d
Luanda (LUAI), Angola 1958 12,000
Addis Ababa (ETH), Ethiopia 1959 6,000
Victoria (SCA), Cameroun 1959 1,400
Monrovia (LIB), Liberia 1960 2,500
Nsukka (UNN), Nigeria 1962 —

754 Proceedings of the Biological Society of Washington

TABLE 12. Oldest and Youngest Herbaria of South America.

Year of Number of
Location Founding Specimens
I. Founded before 1900
1. Rio de Janeiro (RB), Brazil 1808 115,000
2. Buenos Aires (BA), Argentina 1812 80,000
3. Santiago (SGO), Chile 1830 68,742
4. Rio de Janeiro (R), Brazil 1842. 350,000
5. Cordoba (CORD), Argentina 1870 135,000
6. Belem (MG), Brazil 1871 33,500
7. Georgetown (BRG), British Guiana 1879 25,000
8. La Plata (LP), Argentina 1884 220,000
9. Montevideo (MVM), Uruguay 1890 50,000
10. Buenos Aires (BAB), Argentina 1899 140,000
Il. Founded since 1950 (1951-63 )
1. Buenos Aires (IAA), Argentina 1951 600
2. Paraopeba (PMG), Brazil 1951 6,500
3. Cruz Das Almas (IAL), Brazil 1952 8,000
4. Curitiba (IPB), Brazil 1952 5,000
5. Salvador (BAH), Brazil 1952. 2,490
6. Manaus (INPA), Brazil 1954 13,018
7. Porto Alegre (BLA), Brazil 1954 2,500
8. Recife (URM), Brazil 1954 32,000
9. Buenos Aires (BAFC), Argentina 1955 5,000
10. Rio de Janeiro (HB), Brazil 1958 25,000
Tee

Brasilia (UB), Brazil 1963 20,000

759

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Natural history collection symposium 757

TaBLE 14. Comparison of Geographical Distribution of First 100 and
Last 100 Herbarium Foundings.

First 100 Herbaria Last 100 Herbaria

Continent or Region (1545-1840 ) (1947-1963 )1
Africa 0 17
Asia 4 iS
Australia ) 3
British Isles 10 9}
Central America 0 1
Europe 69 13
Malaysia—New Guinea 1 2
Mauritius i 0
Mexico 0 2
North America (excl. Mexico ) 2) 26
Seychelles 0 1
South America $i 15
West Indies 0 2;

1 The ‘Last 100” really number only 99, because to add one more would have re-
quired selecting one from among the 12 founded in different parts of the world in
1946.

TaBLE 15. Frequency Distribution of Herbaria by Size of Collection.

No. Herbaria Percent Total Herbaria
No. Specimens in —_——— Oo
Herbarium Actual Cumulative Actual Cumulative

0-1,000 19 19 2.6 2.6
1,001—25,000 295 314 40.7 43.3
25,001—100,000 202 516 27.9 Tle?
100,001—250,000 102 618 14,1 85.3
250,001—500,000 53 671 The! 92.6
500,001—1,000,000 20 691 2.8 95.4
1,000,001 -2,000,000 20 711 2.8 93:2
2,000,001—3,000,000 6 Ty 0.8 99.0
3,000,001—4,000,000 4 721 0.6 99.6

4,000,00 1—6,500,000 3 724 0.4 100.0

758 Proceedings of the Biological Society of Washington

TABLE 16. Herbaria with 20 or More Staff Members.

Number of Staff

Herbarium Abbreviation Number of
and Location Actual Ideal Specimens

1. MPU, Montpellier, France 46 67 3,150,000
2. SP, Sio Paulo, Brazil 45 y) 85,000
3. K, Kew, England 4] 139 6,500,000
4. LE, Leningrad, USSR 34 128 6,000,000
5. LD, Lund, Sweden 34 32 1,500,000
6. L, Leiden, Netherlands 30 38 1,800,000
7. PC, Paris, France (cryptogams ) 29 26 1,200,000
8. P, Paris, France (phanerogams ) 22 107 5,000,000
9. U, Utrecht, Netherlands Ye 7 350,000
10. BAB, Buenos Aires, Argentina 21 3 140,000
11. US, Washington, D. C., USA 21 64 3,000,000

TOTALS 345 613 28,725,000

1 Probably should be 5,000,000; see footnote, Table 13.

SUMMARY

By DanteL M. CoHEN AND ERNEST A. LACHNER
Bureau of Commercial Fisheries and Smithsonian Institution
Washington, D.C.

The stage for the Symposium was set by Ritterbush, who
placed the topic in broad perspective. He postulated the basis
for the foundation of early collections as naive interest in the
“rare and fabulous,” bound by a stylized and symbolic concept
of natural objects. He made the point that collections had no
scientific value until art and nature were divorced and collec-
tions were acknowledged as examples of the content and diver-
sity of the real world of nature.

The earliest scientific uses of natural history collections still
constitute a prime reason for their existence. To this must be
added, however, the early discovery that patterns of similarities
and differences among organisms could be organized into heir-
archies, which serve as basic documentation for the theory of
evolution. Most speakers proceeded from these two basic as-
sumptions. Even so, Rosewater for mollusks and Duckworth
for entomology pointed out that descriptive and synthetic stud-
ies in which collections are used in traditional ways are far
from complete and that existing resources are by no means com-
mensurate with the uncompleted task. Yochelson and Hotton
emphasized that in the elucidation and comprehension of phy-
logeny, fossil collections are unique in allowing consideration of
the time dimension. The fossil record is sparse and intermittent;
additional collecting is vital. Continued re-evaluation of exist-
ing collections of both fossil and Recent organisms is necessary.

Several speakers described the practical value of collections.
Becklund, in particular, through use of color transparencies in
his presentation, vividly demonstrated the importance of an

57—Proc. Biot. Soc. WasH., Vou. 82, 1969 (759 )

760 Proceedings of the Biological Society of Washington

animal parasite collection. He also described a system of curat-
ing that maintains high standards of documentation and allows
easy access to stored materials. Yochelson discussed the value of
fossils in aging geological formations and Zusi and Duckworth
pointed out the value to agriculture of collection-based studies
on birds and insects. The importance of collections as biologi-
cal standards was treated by Cowan.

Sturtevant reviewed the degree of dependence upon col-
lections of the various branches of anthropology and
concluded that although collections are necessary for some
kinds of research, they are now subordinate to the main objec-
tives of anthropology. He reasoned that over-all, anthropology
can not prosper in a collection-oriented natural history mu-
seum, and that anthropological collections and those who use
them belong in a framework of man-centered studies. He
pointed out, however, many parellel problems in the acquisition
and storage of anthropological and natural history collections.
The plight of anthropological collections is even more serious
than that of plant and animal collections, for even though they
constitute an irreplaceable resource which is potentially the
basis for much important research, there is presently little inter-
est in collection-based research in anthropology. Therefore,
interest is correspondingly small in supplementing and main-
taining collections.

Proceeding under the assumption that collections are neces-
sary for the solution of problems in both basic and applied
science, speakers demonstrated the inadequacy of existing facil-
ities and suggested ways of meeting the increasingly burden-
some problems of housekeeping. Shetler, in particular, pre-
sented hard facts and figures documenting the burden carried
by herbariums. Yochelson reviewed the plight of fossil collec-
tions and made a growth projection.

Few museum administrators or systematists would propose
a halt to collection growth. The problems then are how much
should collections grow and how can they efficiently be used
and maintained?

Several speakers discussed selective collection growth by
group or geographical region and advocated specialization by
museums. We suggest that although this may help efficient

Natural history collection symposium 761

use of collections, it hardly contributes to a solution of the space
problem. Little is gained if the reduction of crowding in one
museum increases it in another. Zusi found that even in a group
as well known as birds, general collections are still important.
He also commented that studies of the comparative biology of
organisms will require not only the maintenance of traditional
kinds of museum collections but also the storage of specimens
prepared in many ways. Data taken from living organisms will
also require systematic storage. In birds for example, photo-
graphs, tapes of songs, and samples of egg whites and blood will
require permanent or temporary curating and storage. Duck-
worth commented on similar problems in insects, particularly
regarding different morphological stages in the insect’s life his-
tory. Modern procedures in taxonomy, combined with greater
ease of data processing, not only allow, but demand the consid-
eration of larger samples than in the past. Then too, as man
modifies the earth with increasing vigor, species once so com-
mon that they were barely worth retaining in collections are
now found only in collections. Specimens that today may be
considered candidates for the trash can could be our only pre-
pollution (chemical-nuclear-thermal) record of a disappearing
environment—a_ biological base line of irreplaceable value.
Sturtevant made analogous comments on collecting anthropo-
logical objects.

Much of the ultimate value of collections lies in what we do
not know about them. Given such a situation, the specialists’
educated guess is our best guide as to what and how much to
save. Regarding our present level of attainment in systematics,
there is no doubt that collection size must not remain on a
plateau based on present holdings. Expand they must!

We suggest that many major collections are not efficiently
housed. Historically, research collections grew in association
with display museums and are located in the high-density pop-
ulation centers of cities, where land costs are high and architec-
tural requirements dictate the construction of elegant, prohibi-
tively expensive structures. Little imagination would be re-
quired to house researchers, libraries and larger collections in
less costly quarters—away from display museums. Divorcing

762 Proceedings of the Biological Society of Washington

the two may sacrifice part of a museum’s integrity, but this is
far less costly than arbitrarily stopping growth of collections.

Good housekeeping was stressed by many speakers as vital
to maintaining access to collections and information about
them. The recognition of professional collection managers was
suggested several times as a step that would relieve researchers
of routine curating. The most widely discussed measure was
electronic data processing (EDP). Shetler viewed it as a signif-
icant innovation in collection and information maintenance on
an international basis. Manning described an EDP project cur-
rently in operation, and Sturtevant noted its use in anthropologi-
cal collections. Zusi questioned the general applicability of
EDP to bird collections but suggested that it could be useful in
some research projects. Becklund and Yochelson felt that cost
mitigates against EDP for their special curatorial needs. Un-
fortunately, no cost figures for EDP were presented. Any
thorough assessment of the future of EDP in the management
of collections must be based on the premise that, although
EDP can supplement collections, it cannot replace them. Also,
the quality of information coming out of a system is limited by
the quality of information fed into it.

In conclusion, we note that to maintain natural history col-
lections adequately is expensive; to neglect them is too costly to
contemplate. The compromise of funding them inadequately
is coming increasingly nearer the latter alternative.

No. 58, pp. 763-766 5 February 1970

PROCEEDINGS
OF THE

BIOLOGICAL SOCIETY OF WASHINGTON

THE IDENTITY OF OLIGODON CYCLURUS (CANTOR,
1839) AND REVALIDATION OF OLIGODON
BREVICAUDA (STEINDACHNER, 1867)
(SERPENTES: COLUBRIDAE)

By Smvwon M. CAamMppen-MAIN
Smithsonian Institution, Washington, D. C.

Simotes brevicauda was described by Steindachner (1867:
61) on the basis of a single specimen lacking precise locality
data from “Cochinchina.” Boulenger (1894:219) placed it
in the synonymy of Simotes cyclurus (Cantor). Since that
time, Steindachner’s brevicauda has appeared in the literature
only as a synonym of Oligodon cyclurus (Cantor). I have
recently examined its holotype, and find that it represents a
valid species of the genus Oligodon. Oligodon brevicauda
(Steindachner) is a secondary homonym of Oligodon brevi-
cauda Ginther (1862:58), and a new name must be proposed.

Oligodon analepticos new name

Simotes brevicauda Steindachner 1867:61.

Holotype: Nat. Mus. Wien 16530 from “Cochinchina,” collected by
Verreaux in 1865.

Diagnosis: Scales smooth, in 19 rows at neck, reducing to 17 rows
between 80th and 97th ventral, reducing to 15 rows between 95th and
117th ventral. Maxillary teeth 10 or 11, last three abruptly enlarged
and recurved. Venter immaculate. fle

Redescription of holotype: Rostral large; visible from above. Parietals
longer than frontal; distance from frontal to rostral less than half length
of frontal. Loreal present. Eight upper labials, fourth and fifth bor-
dering eye. Eye also bordered by presubocular, preocular, supraocular

and two postoculars. Temporals =. + 2. Scales smooth, reducing as

3+4(93 4145(114 :
follows 19 a oa 17 rarer Ten maxillary teeth. Ventrals

162; caudals 44. Twelve dark brown blotches on body, separated by

J ART 1SO58=Proc. Brot. Soc. Wasu., Vou. 82, 1970 (763)
> Wy N
FEB 121970 }
Ba
es

764 Proceedings of the Biological Society of Washington

series of three darker reticulations; two dorsal blotches on tail. Venter
immaculate. Length of head 22.7 mm. Length of tail 71.0 mm. Total
length 620.0 mm. Sex: female.

Additional specimens referred to O. analepticos: Nat. Mus. Wien
19166:1-4, Annam; USNM 95080, 90409, Bao Loc; USNM 146178-79,
Fyan; Nat. Mus. Wien 19167:1, 19167:2, Phuc Son; USNM 144218,
13 km. west Poste M’Drak; USNM 164373, near Bong Son; USNM
163859, near Chu Lai. All localities are in Vietnam.

There has been, and still is, considerable confusion as to the validity
of several taxa now referred to the genus Oligodon. Perhaps the most
confusing are the members of the cyclurus group. This is at least
partially due to the inadequate original description of Cantor (1839:50),
in which he described Coronella cyclura but failed to designate a type
locality. Subsequently the holotype of cyclura has been lost, leaving
very little on which to form an opinion as to the identity of Cantor’s
species. In the interest of nomenclatural stability, it seems best to
designate a neotype, thus establishing a type locality in the area
where most specimens that recent workers have assigned to cyclurus
have been collected. I, therefore, designate as the neotype USNM
72067, a male collected in Bangkok, Thailand, by Hugh M. Smith on
September 8, 1934.

Oligodon cyclurus (Cantor )

Coronella cyclura Cantor, 1839:50.
Oligodon cyclurus-Smith, 1943:202.

Neotype: USNM 72067 from Bangkok, Thailand, collected by H. M.
Smith, September 8, 1934.

Diagnosis: Scales smooth, in 21 rows at neck, reducing to 19 rows
between 95th and 113th ventral, reducing to 17 rows between 108th
and 152nd ventral. Maxillary teeth 10, rarely 9 or 11, last three
abruptly enlarged and recurved. Venter immaculate.

Description of neotype: Rostral large, visible from above. Parietals
longer than frontal; distance from frontal to rostral less than half length
of frontal. Loreal present. Eight upper labials, fourth and fifth
bordering eye. Eye also bordered by presubocular, preocular, supra-

ocular, and two postoculars. Temporals 2 + > Scales smooth reduc-

ing as follows 214+5/(105)194+5(111) 17, Ten maxillary teeth. Ven-
44+5(105) 445(112)

trals 162; caudals 44. Fourteen dark brown dorsal blotches on body,
separated by series of three darker reticulations; four dorsal blotches on
tail. Venter immaculate. Length of head 17.5 mm. Length of tail 71.0
mm. Total length 456.0 mm. Sex: male.

Additional specimens referred to O. cyclurus:; USNM 70324, 70326,
72066-72069, 75683, 75684, 76122, 79472-79474, 81838, 83432, 94757,
94758, 100994, 101287, Bangkok; USNM 70355, 101290, Lam Tong
Lam; USNM 94931, Sam Roi Yot; USNM 76090, Pichit. All localities
are in Thailand.

Oligodon cyclurus and brevicauda 765

Oligodon analepticos may easily be distinguished from O. cyclurus by
both geography and by dorsal scale row count. In all specimens exam-
ined from Thailand and Laos the dorsals reduce from 21 to 19 then to 17,
as opposed to all South Vietnamese specimens which reduce from 19
to 17 then to 15 scale rows. On the basis of material examined sym-
patry can not be demonstrated.

I am indebted to Dr. Joseph Eiselt ( Naturhistorisches Museum, Wien,
Austria) for the loan of the holotype of Simotes brevicauda Stein-
dachner, and other material. Drs. J. A. Peters and G. Zug reviewed
and commented on the manuscript.

LITERATURE CITED

BouLencer, G. A. 1894. Catalogue of the snakes in the British Mu-
seum (Natural History). Vol. 2. London.

Cantor, T. 1839. Spicilegium Serpentium Indicorum. Proc. Zool.
Soc. London. 1839(7): 31-34, 49-55.

GUnTueErR, A. 1862. On new species of snakes in the collection of
the British Museum. Ann. Mag. Nat. Hist. Ser. 3, 9: 52-59.

STEINDACHNER, F. 1867. Reise der Osterreichischen Fregatte Novara
um die Erde in den Jahren 1857, 1858. Zoologischer Theil.
Bd. 2, Reptilien. Wien.

766 Proceedings of the Biological Society of Washington

Vol 767-776 5 February 1970

PROCEEDINGS
OF THE

BIOLOGICAL SOCIETY OF WASHINGTON

DESCRIPTIONS OF ADDITIONAL FORMS OF BIRDS
FROM PANAMA AND COLOMBIA

By ALEXANDER WETMORE
Smithsonian Institution, Washington, D.C.

The descriptions and other information presented in the
following pages have come to my attention during contin-
uing studies of the collections of birds from Panama and
Colombia in the U.S. National Museum. They include com-
parisons made in collections in other institutions, in addition
to those available in Washington.

Metallura primolinus recisa new subspecies

Characters: Similar to Metallura primolinus primolinus Bourcier,' but
with bill shorter and more slender; slightly duller green on dorsal surface;
male with dark throat patch smaller, narrower.

Description: Type, USNM 436,301, ¢, from Paramo de Frontino, 3600
meters elevation, Antioquia, Colombia, collected by M. A. Carriker,
Jr., 29 August 1951 (original number 21030). Upper surface from
the forehead to the upper tail coverts, including the lesser wing coverts,
metallic dark cress green; upper surface of tail dull metallic green; pri-
maries, secondaries and greater wing coverts dull purplish black; middle
wing coverts faintly dull bronze; feathers in front of eye tipped with
rufous; under surface, including under wing coverts, like back, but
faintly more yellowish green; a narrow, elongated gorget, extending
from the chin down the center of the foreneck, viewed by direct illumi-
nation, shining light metallic green; seen from the opposite angle, dull
black, the individual feathers tipped and edged lightly with cinnamon;
center of abdomen barred centrally with dull green, tipped narrowly
with dull cinnamon-buff, producing a mixed pattern; a prominent white
tibial tuft on either side; under tail coverts basally dark metallic green,
tipped with cinnamon; edge of wing lined narrowly with cinnamon;
under surface of tail bright metallic green.

Female, foreneck pale cinnamon, with the individual feathers tipped

1 Metallura primolinus Bourcier, Rev. et Mag. Zool., ser. 2, vol. 5, July, 1853,

ei Wie ity of Laguano, Napo, Ecuador. )
OWN if
NA Brot. Soc, Wass, Vor. 82, 1970 (767)

FEB 12 970)
}
SiBRARIED~

768 Proceedings of the Biological Society of Washington

with a spot of metallic dark green; breast similar, but with the base
color pale cinnamon, varying to white, and the green spots much
larger, in some areas covering the base color; otherwise like the male.

Immature male, with no gorget, and with very slight indication of
the markings of the female on foreneck and breast.

Measurements: Males (8 specimens), wing 54.9-60.2 (58.2), tail
36.2-40.7 (39.2), culmen from base 14.2-15.2 (14.8) mm.

Females (7 specimens), wing 54.0-57.0 (55.0), tail 35.1-37.9 (36.7),
culmen from base 13.8-15.0 (14.7) mm.

Range: The higher levels of the Paramo de Frontino, in the Cordillera
Occidental, western Antioquia, Colombia.

Remarks: Typical Metallura primolinus primolinus is known mainly
from specimens taken in the Andean chain in northern Ecuador, moun-
tains which in effect continue directly north through Colombia as the
Cordillera Central. The record for the species in the Paramo de Frontino
is the most northern at present, and is in the western division of the
Cordillera. It seems probable that M. p. recisa is confined to the
latter range.

Peters (1945, p. 119) lists primolinus as a race of williami, a species
of similar general appearance but with the tail wholly deep blue, being
brilliant shining blue on the under surface. The entirely green tail of
the populations of primolinus are so definitely distinct in the specimens
seen that it is appropriate to treat it as a separate species.

It should be noted that Bourcier named Metallura primolinus in
honor of the son of Count Primoli, grandson of Prince Charles Lucien
Bonaparte. As the word is of masculine gender it stands in apposition
to the generic name.

Measurements of typical M. p. primolinus from Ecuador are as follows:
Males (4 specimens), wing 55.4-59.9 (58.5), tail 36.6-39.5 (38.4),
culmen from base 16.3-17.4 (17.0) mm.

Females (11 specimens), wing 55.3-58.0 (56.7), tail 35.0-39.6 (36.9),
culmen from base 15.8-17.5 (16.8) mm.

Etymology: The subspecific name for this race is from the Latin
adjective recisus, meaning short, in reference to the bill.

Specimens examined in comparisons: Metallura primelinus primolinus,
Ecuador—above Baeza, 6, 3 9; Papallacta, 5 ¢,7 9.

Sittasomus griseicapillus enochrus new subspecies

Characters: Similar to Sittasomus griseicapillus veraguensis Aldrich,”
but with the under surface from throat to abdomen paler, more greenish;
under tail coverts lighter, more cinnamon-brown; crown, hindneck and
back paler, faintly more greenish gray; rump and upper tail coverts
lighter cinnamon-brown.

2 Sittasomus griseicapillus veraguensis Aldrich, Sci. Publ. Cleveland Mus. Nat.
Hist., vol. 7, 31 August, 1937, p. 83. (Rio Mariato, 16 kilometers east of Golfo
de Montijo, Veraguas, Panama. )

New subspecies of birds 769

Description: Holotype, USNM 411264, ¢, from 6 kilometers north of
Colosé, northern Department of Bolivar, Colombia, collected 20 October
1948, by M. A. Carriker, Jr. (original number 14127). Crown and
hindneck dark grayish olive; back and scapulars dull buffy brown;
rump and upper tail coverts tawny; lesser and middle wing coverts
hair brown; greater coverts and primary coverts with inner webs
chaetura drab, outer webs drab; secondaries externally between russet
and tawny; inner webs distally fuscous-black except at tip; primaries
fuscous-black, with the outer webs dull ochraceous-tawny; foreneck and
breast light grayish olive; abdomen washed lightly with deep olive-buff;
under tail coverts ochraceous-tawny; inner under wing coverts very pale
buffy white, changing externally to grayish olive; inner webs of second-
aries and primaries toward base light ochraceous-buff.

Measurements: Males (3 specimens), wing 75.6-81.0 (79.1), tail
71.8-75.0 (73.2), culmen from base 16.0-16.2 (16.1), tarsus 16.1-16.8
(16.5) mm.

Female (1 specimen), wing 72.3, tail 67.5, culmen from base 15.8,
tarsus 16.2 mm.

Remarks: As a species Sittasomus griseicapillus, the Olivaceous
Woodcreeper, has been little known in northern Colombia. Currently,
the few records have been listed under the subspecies name levis,
which is the race now confined to the Province of Chiriqui in western
Panama, with S. g. veraguensis covering the area from eastern Veraguas
east through Darién. As I have found veraguensis in Darién on Cerro
Mali, a spur of Cerro Tacarcuna on the border between Panama and
Colombia, it is probable that veraguensis extends into the latter country.

In addition, in the series of specimens collected by M. A. Carriker,
Jr., I find that, as listed beyond, S. g. tachirensis Phelps and Phelps, Jr.,
named from Tachira in western Venezuela, ranges west to the central
Magdalena valley in Colombia. Further, the somewhat darker popu-
lation S. g. perijanus Phelps and Gilliard, described from the Venezuelan
side of the Sierra de Perijé, extends across to the Colombian side of this
mountain range in northwestern Magdalena and western Guajira.

It is probable that the race enochrus occupies the area from northern
Bolivar west through northern Cérdoba. In size it is like veraguensis, in
which the wing in males ranges from 74.0 to 80.4, with an average of
78.0 mm. The other two here recorded for Colombia are slightly larger,
as shown by the following measurements of males:

Sittasomus g. tachirensis (8 specimens), wing 80.2-86.8 (83.6) mm.
Sittasomus g. perijanus (5 specimens), wing 78.6—87.7 (82.5) mm.

Etymology: The name for this race has been taken from the Greek
enochros, meaning rather pale.

Specimens examined in comparisons: Sittasomus griseicapillus levis,
Panam&—Chiriqui: Sereno, ?; Santa Clara, 2 ¢; El Volcan, 8 ¢, 109;
Boquete, 3 6, 92.

Sittasomus griseicapillus veraguensis, Panama—Los Santos: Los Asi-
entos, 6, 9. Herrera: Parita, ¢; Portobellilo, 9. Province of Panama:

770 Proceedings of the Biological Society of Washington

La Campana, 6; Utivé, 6; Pacora, 6; Chepo, 6, 2; Chiman, 6. Darién:
Cerro Mali, 6, 9; Cana, 6, @.

Sittasomus griseicapillus tachirensis, Colombia—Caldas: Hacienda
Sofia, 2 ¢, 9. Antioquia: La Bodega, 6, @. Bolivar: Volador, 2 ¢.
Norte de Santander: Convencién, 2 9; Guamalito, ¢, 2; Buenos
Aires, 3 6.

Glyphorhynchus spirurus pallidulus new subspecies

Characters: Similar to Glyphorhynchus spirurus sublestus Peters,? but
paler, more grayish olive on lower surface; above, paler, less reddish
brown; rump and upper tail coverts brighter, lighter cinnamon-rufous;
chin and upper throat slightly paler buff.

Description: Holotype, USNM 423458, é, from Charco del Toro, Rio
Majé, Province of Panama, Panama, collected 28 March 1950, by A.
Wetmore and W. M. Perrygo (original number 16004). Forehead,
adjacent to base of bill, very narrowly pale gray; crown olive-brown;
forehead with very narrow, faint shaft lines of pale buff; hindneck, back,
scapulars and wing coverts between bister and snuff brown; alula
Saccardo’s umber; primary coverts dull black, with outer webs between
bister and snuff brown; rump and upper tail coverts between tawny and
russet; secondaries russet; primaries deep olive, with outer webs cin-
namon-brown; tail russet, with shafts ferruginous; narrow superciliary
streak dull buff; side of head behind eye olive-brown, the auricular
area streaked lightly with dull gray; chin and upper throat pinkish buff,
spotted lightly with olive-brown; lower foreneck, uppermost breast, and
side of neck somewhat dark buffy brown, with lanceolate shaft lines of
buffy white, narrower laterally, broader at center; lower breast, sides
and abdomen grayish brown (between buffy brown and citrine-drab )
with narrow dull white shaft lines on breast; under tail coverts Dresden
brown, with shaft lines of dull white; axillars and under wing coverts
white, with a spot of sepia on outer side near center; a band of pale
cinnamon-buff across the inner webs of the secondaries and inner pri-
maries.

Measurements: Males (17 from eastern provinces of Panama and
Colén, Darién and San Blas), wing 71.7-75.5 (73.9), tail 62.8-69.9
(66.6), culmen from base 12.3-14.2 (13.0), tarsus 16.5-17.8 (17.2) mm.

Females (20 from eastern Province of Panama, Darién and San Blas),
wing 64.6-73.8 (69.5), tail 59.0-68.0 (62.6), culmen from base 12.2—
13.5 (12.9), tarsus 16.0-17.7 (16.7) mm.

Holotype, male, wing 74.2, tail 67.4, culmen from base 14.2, tarsus
17.5 mm.

Range: Tropical lowlands; on the Pacific slope through eastern
Province of Panama from the Cerro Azul through Darién (except the
southwest on the Rio Jaqué); on the Caribbean side in eastern Coldén,
including the Chagres Valley back of Madden Lake, and San Blas,

8 Glyphorhynchus spirurus sublestus Peters, Bull. Mus. Comp. Zoél., vol. 69,
October 1929, p. 443. (Changuinola, Bocas del Toro, Panama.)

New subspecies of birds 771

extending in Colombia to the extreme northern tip of Chocd (Acandi,
Rio Cuti, eastern slope of Cerro Tacarcuna); ranging to 1450 meters in
mountain areas.

Remarks: The Wedge-billed Woodcreeper is the most abundant spe-
cies of its family through the Isthmus of Panama, found in the more
humid areas wherever there is forest cover. Three populations, differing
slightly in color, inhabit the region.

Glyphorhynchus spirurus sublestus Peters, darker, more reddish brown
above, darker on the lower surface, with the throat more cinnamon-
buff, ranges through the lower levels of Costa Rica and western Panama,
on the western slope to the base of the volcano in Chiriqui, on the Carib-
bean side through the northern Canal Zone to Cerro Bruja in eastern
Colén, where it intergrades with the paler pallidulus. At the eastern
end of its range in northwestern Colombia the race pallidulus merges
with another darker race Glyphorhynchus spirurus subrufescens, which
is more olive, less reddish brown above, and more olive on the ventral
surface than the race sublestus. It also is slightly smaller, with the
wing in males 66.1 to 71.8 (average 69.0) mm., and in females 63.5: to
69.5 (average 66.2) mm. This form comes into southeastern Darién in
the valley of the Rio Jaqué. From there it ranges south along the Pacific
slope of Colombia to the Department of Narifio, probably to north-
western Ecuador. Eastward it extends through the lower Atrato valley
in northern Antioquia (Villa Artiaga) and northern Chocd. Another
variant appears in the Sint Valley in the Department of Cérdoba and
eastern Antioquia, recognized as Glyphorhynchus spirurus integratus.
This group, paler above and below than subrufescens, with the throat
paler buff (but darker than pallidulus), ranges through the departments
of Bolivar to Santander and Norte de Santander into western Venezuela.

Etymology: The subspecific name for the race described above has
been taken from the Latin adjective pallidulus, somewhat pale, from
its lighter colors.

Specimens examined in comparisons: Glyphorhynchus spirurus sub-
lestus, Costa Rica—La Vijagua, 3 ¢,3 2; Pozo Azul, ¢; Rio Matina, ¢;
Bonilla, 2 9; Buenos Aires, 9; Pacuare, 9; Jiménez, ¢; Reventazén,
6, 9; Talamanca, ¢. Panama—Chiriqui: El Volcan, 2 ¢; Santa
Clara, 2; Puerto Armuelles, 2. Coclé: Rio Guabal, ¢; El Uracillo, ¢.
Canal Zone: Gatun, 3 ¢, 3 2; Lion Hill, ¢. Colén: Cerro Bruja, 2 @.

Glyphorhynchus spirurus subrufescens, Panama—Darién: Jaqué, 9;
Rio Jaqué, 2 ¢@. Colombia—Antioquia: Hacienda Potreros, ¢; Villa
Artiaga, 6 6,3 9. Chocéd: Rio Jurubida, 2 ¢, 4 9; Rio Nuqui, ¢,4 9.
Valle: Punto Muchimbo, ¢. Narifo: La Guayacana, 2 ¢, 2 9. Coéor-
doba: Socarré (Rio Sint), ¢, 2; Quebrada Salvajin (Rio Sind), 6, 9.

Glyphorhynchus spirurus integratus, Colombia—Antioquia: El Pes-
cado, ¢, 9; Valdivia, 4; Hacienda Belén, 3 6; El Real, 2¢; La Raya,
6 6,5 9; Regeneracién, 2 6. Bolivar: Volador, 2. Santander: Haci-
enda Santana, 6 6,5 2. Norte de Santander: Petrélea, 9; Bellavista,
26.

772 Proceedings of the Biological Society of Washington

Xenops rutilans incomptus new subspecies

Characters: Similar to Xenops rutilans septentrionalis Zimmer,* but
smaller; pale streaks on lower surface, crown and back narrower; rump
and upper tail coverts faintly darker.

Description: Holotype, Museum of Comparative Zoology No. 140709,
3, from Cana, Cerro Pirre, Darién, Panama, collected 31 July 1928, by
R. R. Benson (original number 460). Crown and hindneck sooty brown,
streaked narrowly with pale brownish buff; back and scapulars dull
cinnamon-brown, with the upper back streaked narrowly with cinnamon-
buff; rump, upper tail coverts and tail cinnamon-rufous; inner web of
third and fourth rectrices from outside black; fifth rectrix lined centrally
on inner web with dusky; wing coverts with outer webs like back, inner
webs dusky; secondaries cinnamon-rufous with a concealed distal spot
of black; primaries with inner webs black, outer webs cinnamon-rufous;
lores, and a streak from center of eye back along side of crown, dull
white; side of head dusky lined with dull white; a short, narrow streak
of slightly elongated feathers on the lower margin of the side of the
head at the back clear white; chin, throat and upper foreneck clear white;
under surface pale grayish brown, washed faintly with cinnamon on
lower abdomen and flanks; under tail coverts pale dull cinnamon; sides
of foreneck, breast, upper abdomen and under tail coverts lined narrowly
with dull white; axillars white; edge of wing, and inner under wing
coverts dull white; rest cinnamon-buff.

Measurements: Male (holotype), wing 62.0, tail 41.2, culmen from
base 12.9, tarsus 14.7 mm.

Female (one specimen), wing 62.7, tail 41.9, culmen from base 13.0,
tarsus 15.0 mm.

Remarks: The two specimens on which this form is based, reported
originally by Ludlow Griscom (1929, p. 171), were referred pro-
visionally to the race heterurus of northern Colombia, but with recog-
nition that they appeared to differ from the scanty material then available
for comparison. From the series of heterurus now at hand they differ in
smaller size, reduced streaking, definitely grayer under surface, and deeper
cinnamon hue above, especially on the rump and upper tail coverts.

Etymology: The subspecific name is from the Latin adjective in-
comptus, in the sense of unadorned, from the reduction in markings
compared to those of its near relatives.

Specimens examined in comparisons: Xenops rutilans septentrionalis,
Panama—Chiriqui, 4 6, 4 9.

Xenops rutilans heterurus, Colombia—Antioquia: Hacienda Zulaiba,
$, 9%; Hacienda La Ilusién, Rio Urrao, @; Hacienda Potreros, 9@.
Cauca: Hacienda La Capilla, ¢, 2 ?; Tijeras, Moscopan, 9. Huila:
La Candela, 2 ¢; Belén, 3 ¢. Santander: Hacienda Las Vegas, 3 @.

4 Xenops rutilans septentrionalis Zimmer, Proc. Biol. Soc. Washington, vol. 42,
March 25, 1929, p. 82. (Guayabo, Costa Rica. )

New subspecies of birds 773

Thamnophilus doliatus nesiotes new subspecies

Characters: Similar to Thamnophilus doliatus nigricristatus Lawrence,°
but male darker; under surface with black bars heavier, and the white
interspaces correspondingly reduced; female also darker; slightly larger.

Description: Holotype, USNM 471358, ¢, from Rio Cacique, Isla del
Rey, Archipiélago de las Perlas, Panama, collected 27 January 1960, by
Alexander Wetmore (original no. 23186). Forehead pale grayish white,
streaked lightly with black; crown black, tipped lightly with white at
the back (these paler markings partly concealed); hindneck black,
streaked narrowly with white; back and scapulars black, barred with
white (the light bars one-half or less as wide as the black ones); wings
black, barred and spotted narrowly with white; tail black, marked
narrowly with white on the outer edge of both webs; side of head black,
lined with pale grayish white; throat and upper foreneck white,
streaked rather narrowly with black; rest of under surface, including
sides, flanks and under tail coverts, white barred with black, the
barring narrower on the abdomen which thus appears whiter; under
wing coverts white spotted lightly with black; inner webs of flight
feathers barred widely with white.

Measurements: Males (11 from islas del Rey, Cafias and Pedro Gon-
zalez), wing 70.3-74.8 (72.8), tail 53.8-58.8 (56.2), culmen from base
20.2-22..4 (21.4), tarsus 26.2-27.9 (27.1) mm.

Females (7 from islas del Rey and Pedro Gonzalez), wing 69.4-73.6
(71.4), tail 53.4-57.7 (56.3), culmen from base 20.2-22.4 (21.4),
tarsus 26.2-27.9 (27.1) mm.

Holotype, male, wing 72.5, tail 55.7, culmen from base 22.3, tarsus 27.5
mm.

Range: Archipiélago de las Perlas, Gulf of Panama, Panama, where
recorded from islas Pedro Gonzalez, del Rey, Viveros and Cafias.

Remarks: In field work in the archipelago I was interested to find
that the Barred Antshrike was not present on Isla San José which is one
of the larger land masses in the group, but isolated to the southwest, nor
have I encountered it on the small islands Contadora, Saboga, Chapera,
Santelmo and Bayoneta. Possibly lack of water supply may be a factor
on these smaller islands as both Viveros and Canas, where the bird is
present, while small, have permanent springs.

Etymology: The name for this insular population is from the Greek
nesiotes, an islander.

Specimens examined in comparisons: Thamnophilus doliatus pacificus,
Nicaragua—Sucuya, 2 ¢, 9; Chinandega, ¢ (type). Costa Rica—La
Palma, ¢; Bebedero, 2 6, 2; Pozo Azul, 9; Bolsdn, 4 6,3 9; El Gen-
eral, 2 6, 2. Panama—Chiriqui: Divala, ¢, 2 9; Concepcidn, $, 9°.

Thamnophilus doliatus nigricristatus, Panama—Chiriqui: San Félix, 9;

5 Thamnophilus nigricristatus Lawrence, Proc. Acad. Nat. Sci. Philadelphia,
1865, p. 107. (Lion Hill Station, Panama Railroad, Atlantic slope, Canal Zone,
Panama. )

774 Proceedings of the Biological Society of Washington

Las Lajas, 2 ¢; Quebrada Piedra, 9. Veraguas: Sona, 7 6, 4 9; Rio
de Jestis, ¢; Chitra, 9. Coclé: Aguadulce, 2; El Copé, ¢; El Potrero,
24,5 @; Gago, 2 6, 9; El Uracillo, $, 9. Los Santos: Las Palmitas,
2 46,3 9; Ensenado Venado, 6,2 9; Punta Mala, ¢; Pedasi, 2 6,3 9;
Tonosi, 2 6, 2 9; Los Santos, ¢. Herrera: Parita, 4 6, 3 9; La
Cabuya, 2; El] Barrero, 6, 2. Western Province of Panama: Nueva
Gorgona, 3 6, 9; Cerro Chame, ¢. Canal Zone: Farfan, 9; Fort Clay-
ton, 6, 9; Corozal, ¢, 9; Pedro Miguel, ¢; Miraflores, 9; Tabernilla,
24,2 9; Frijoles, ¢, 2; Bas Obispo, 6, 2; Rio Indio, ¢; Lion Hill, 4;
Juan Mina, 2 ¢, 4 9. Eastern Province of Panama: Panama, 2 ¢, 9;
Rio Abajo, 2; Pacora, 2 ¢, 2 9; Chico, ¢, 2 9; Chepo, 246; El Llano,
9; Canita, ¢. Colombia—Antioquia: Necocli, ¢, 9. Cérdoba: Tierra
Alta, 2 ¢, 2 9; Socarré, 6; Pueblo Nuevo, 2 6, @. Bolivar: Simiti,
3 4; Santa Rosa, 6, 9; Rio Viejo, 2 4, 9; Colosé, ¢. Magdalena:
La Gloria, ¢; Hacienda La Esperanza, 2 ¢, 9; Codazzi, 6. Guajira:
La Cueva, 3 6; Los Gorros, ¢, 9.

Oryzoborus crassirostris loftini new subspecies

Characters: Similar to Oryzoborus crassirostris nuttingi Ridgway,®
but female distinctly darker, less rufescent brown on both upper and
lower surface; slightly smaller, with bill somewhat more slender.

Description: Holotype, USNM 533762, 9°, from Almirante, Bocas del
Toro, Panama, collected by P. Kirmse and T. V. Heatley, 14 October
1967. Crown, hindneck, back and scapulars olive-brown, changing to
bister on the lower rump and upper tail coverts; lesser wing coverts bister;
wings otherwise dull fuscous-black, with the middle and greater coverts,
secondaries and inner primaries edged with dull bister; tail fuscous-black;
side of head somewhat duller than bister; under surface slightly darker
than snuff-brown; flanks and under coverts bister; (a single aberrant
pure white, albinistic feather on the center of the right side); axillars and
outer wing coverts olive-brown; inner under wing coverts and edgings
of primaries on underside dull grayish white. Bill, in life, very dark
brown.

Measurements: Males (2 from Almirante), wing 67.2-67.4 (67.3),
tail 58.7-60.7 (59.7), culmen from base 18.0-18.3 (18.1), transverse
width of mandible at base 13.3-14.0 (13.6), tarsus 18.2-18.8 (18.5) mm.

Female (holotype, from Almirante), wing 65.0, tail 59.0, culmen
from base 18.1, width of mandible at base 13.7, tarsus 18.2 mm.

Range: Known from Almirante, Bocas del Toro, near the western end
of the Caribbean slope, Panama.

Remarks: The first intimation of the presence of this small, large-
billed finch in Panama was a male found at Almirante 10 June 1965,
caught in the edge of a closed mist net set by men capturing birds for
banding under the direction of Dr. Pedro Galindo. As it was partly

6 Oryzoborus nuttingi Ridgway, Proc. U.S. Nat. Mus., vol. 6, April 26, 1884,
p. 401. (Hacienda Los Sabalos, Rfo San Juan, Chontales, Nicaragua. )

New subspecies of birds 775

decomposed, it could be preserved only in part as a flattened specimen.
Those working with birds in the area were alerted to watch for the
species, but others were not found until two years later. Two males and
a female were netted on 13, 14 and 30 October 1967 by P. Kirmse, V. M.
Kleen and T. V. Heatley, operating the banding station under the direc-
tion of Dr. Horace Loftin. The species has been known previously
in Central America from the Caribbean slope in eastern Nicaragua and
northern Costa Rica. In South America it is found from northwestern
Colombia, Venezuela and Trinidad to western Ecuador, eastern Pert
and southern Brazil. The South American populations are distinct in
having white or partly white axillars and under wing coverts, and a
small white wing speculum (varying in size) in males. In some males
there is concealed white also on the base of the rectrices.

Measurements of Oryzoborus crassirostris nuttingi are as follows:
Males (11 from Nicaragua, 1 from Costa Rica), wing 69.0-71.0 (69.8),
tail 61.0-66.3 (62.9), culmen from base 18.0-19.1 (18.6), transverse
width of mandible at base 13.9-15.7 (14.7), tarsus 18.3-20.1 (19.1) mm.

Females (5 specimens), wing 66.0-69.1 (67.2), tail 59.4-65.4 (60.4),
culmen from base 18.0-19.7 (18.7), transverse width of mandible at
base 13.7-14.6 (14.3), tarsus 14.3-15.9 (14.7) mm.

Etymology: This addition to the avifauna of Panama is named for
Dr. Horace Loftin, head of the Florida State University Center for
Tropical Studies in the Canal Zone, in recognition of his interest in the
fauna of the Republic.

Specimens examined in comparisons: Oryzoborus crassirostris nuttingi,
Nicaragua—Greytown, 3 ¢, 2; Rio Escondido, 9; Los Sabalos, Rio San
Juan, 7 2.5 9.

Oryzoborus crassirostris crassirostris, Colombia—Cérdoba: Tierra Alta,
3; Socarré, 9; Quebrada Salvajin, 2 ¢ im., 4 9. Antioquia: Taraza, ?;
Hacienda Belén, ¢. Bolivar: La Raya, 4 6, @.

ADDITIONS TO THE List or Birps RECORDED FROM COLOMBIA

The following, based on collections in the U.S. National Museum,
made in Colombia by M. A. Carriker, Jr., add to the list as published
by R. Meyer de Schauensee in the second printing of his volume, The
Birds of Colombia and adjacent areas of South and Central America,
published originally in 1964.

Racquet-tailed Hummingbird, Ocreatus underwoodi discifer Heine.

Carriker collected two adult, two immature males, and a female in
forest between 1675 and 1980 meters elevation on the Sierra de Perija,
above Hiroca, Magdalena, from 17 April to 1 May 1942. He found
the species again near Palo Gordo, Norte de Santander, where he
secured two males on 14 and 22 November 1947. One of these was
taken in the shade trees over coffee, the other in heavy forest. The race,
described from the mountains near Mérida, Venezuela, ranges in that
country to the intermediate levels on the eastern face of the Sierra de
Perijaé. It is no surprise therefore to find that it occurs also on the

776 Proceedings of the Biological Society of Washington

Colombian slopes of the range. Males are similar to nominate under-
woodi. The females differ from that race in having the chin and center
of throat plain white, with the spotting restricted or absent also on the
center of the breast. In underwoodi these areas are definitely spotted.

Striped-breasted Spinetail, Synallaxis cinnamomea aveledoi
Phelps and Phelps, Jr.

As an addition to the races of this species in Colombia, Carriker
collected two males and three females of aveledoi at Palo Gordo, Norte
de Santander, from 14 to 23 November 1947. These are the first records
for Colombia, theiz identity checked by comparison with the type in
the American Museum of Natural History. The race, found from
northwestern Zulia to Mérida and Tachira in adjacent western Venezuela,
is plain warm brown above, less reddish than any of the other races, and
also is more buffy, less cinnamon on the lower surface.

LITERATURE CITED

Griscom, L. 1929. A collection of birds from Cana, Darien. Bull.
Mus. Comp. Zool., 69(8): 149-190.
Peters, J. L. 1945. Check-list of Birds of the World, 5: 119.

a — ———————

Ls i ee
ra L Varo

wat pe tet

pp. 777-788 5 February 1970

PROCEEDINGS
OF THE

BIOLOGICAL SOCIETY OF WASHINGTON

A NEW SPECIES OF LARGE DIPLOGLOSSUS
(SAURIA: ANGUIDAE) FROM HISPANIOLA

By ALBERT SCHWARTZ
Dept. of Biology, Miami-Dade Junior College,
Miami, Florida

The anguid lizards of the genus Diploglossus Wiegmann
are widespread on the islands of the Greater Antilles. The
number of species on each island, however, varies; Cuba has
but a single species (delasagra Cocteau), and Puerto Rico
likewise has but one galliwasp (pleii Duméril and Bibron).
Jamaica and Hispaniola have a diversity of forms; the former
island has (or had) six species (occiduus Shaw, barbouri
Grant, crusculus Garman, duquesneyi Grant, hewardi Gray,
microblepharis Underwood) and Hispaniola has five species
(costatus Cope, curtissi Grant, darlingtoni Cochran, sep-
soides Gray, stenurus Cope). Another species (montisserrati
Underwood) is the sole representative of this genus in the
Lesser Antilles where it occurs on Montserrat. In addition,
D. costatus occurs on Navassa (between Hispaniola and Ja-
maica) and D. crusculus occurs on the Lesser Cayman Islands
(Little Cayman and Cayman Brac).

Of these species, the least known is D. occiduus. Grant
(1940:109) was not convinced that D. occiduus was extinct,
but Cousens (1956:1) stated that this giant galliwasp had not
been collected in over 100 years. Although D. occiduus had
in early times been reported to live in swamps and to eat fish
and fruit (and thus in its habits and habitats it may have tat
less conspicuous than other of its Jamaican congeners), it:
seems highly unlikely that the species still persists in eee 2
I know of only three specimens in American collections; I=
have examined these lizards in the collection of the Museum— o
of Zoology at the University of Michigan and the Museum |

60—Proc. Brot. Soc. Wasu., Vou. 82, 1970 HET),

778 Proceedings of the Biological Society of Washington

of Comparative Zoology at Harvard University through the
courtesy of Charles F. Walker and Emest E. Williams. If
for no other reason, D. occiduus is the most distinctive of the
Antillean Diploglossus because of its very great size.

When I reviewed the galliwasps of the Hispaniolan costatus
complex, I examined a specimen in the United States National
Museum from Rivicére Bar (= Riviére des Barres) in northern
Haiti; this specimen was collected by W. L. Abbott in 1917
and had been assigned by Cochran (1941:250) to her all-
inclusive “Celestus costatus.” In turn, in my review of this
complex, I consider the lizard as D. stenurus rugosus (1964:
17) and commented upon its very large size (snout—vent length
230 mm) which far exceeded that of any other specimen of
D. stenurus studied. The lizard is badly crushed about the
head, and details of head scutellation (aside from the position
of the angular subocular on both sides) cannot be determined.
I had suspected that possibly this lizard represented still
another distinct taxon from northern Haiti, a taxon related
to the costatus complex (which includes costatus, stenurus
and curtissi), but the condition of the single specimen, its
faded pattern due to length of time in preservative, and the
fact that there was but one animal made its description an
improvident course.

Through the efforts of C. Rhea Warren, herpetological col-
lections were made on Ile de la Tortue off the northern Haitian
coast in 1968. The herpetofauna of this island has been very
poorly known, and through the efforts of Mr. Warren and local
natives, excellent collections including several unexpected
forms have now been collected there. Among the lizards
secured are three species of Diploglossus: costatus, curtissi,
and two specimens of the giant form previously known from
Riviere des Barres. The absence of D. stenurus or immature
representatives of the giant species is remarkable, although
neither costatus nor curtissi is apparently common in the vi-
cinity of Palmiste, whence the recent collections have come. It is
moreover remarkable that the Riviere des Barres lies on the
Haitian coast immediately opposite Ile de la Tortue and just to
the east of the town of St. Louis du Nord. The entire northern
Haitian coast has been poorly represented by D. stenurus

A new Diploglossus 779

(despite many collections from Cap-Haitien and its vicinity in
recent years); this species has been reported from Bombard-
opolis on the Presqu’ile du Nord Ouest on the west, and near
Limbé and near Limonade to the east of the Riviere des
Barres (one specimen from each of these three localities;
Schwartz, 1964:18). However, additional material more re-
cently collected by Richard Thomas shows that D. stenurus is
fairly common near Limbé, and in the vicinity of Limonade,
and inland from Anse 4 Margot, all settlements on the northern
Haitian versant. Thomas’s material is both crucial and critical
as far as comparison of D. stenurus and the new species is
concerned.

Examination of Thomas’s excellent series of 26 D. stenurus
from these localities shows that there is nothing distinctive in
size as far as they are concerned. The largest specimen
(Albert Schwartz Field Series [ASFS] V9957) is a male with a
snout—vent length of 138 mm; this measurement lies within the
known extremes of male D. stenurus (maximum size 172 mm
snout—vent length in D. s. stenurus, the largest of the four sub-
species). Patterned D. stenurus from northern Haiti have
the back covered with closely appressed dark brown and
fragmented herringbones; the largest D. stenurus from this
region (ASFS V9957) shows a dissolution of this pattern to one
of dark longitudinal lines, the lines lying on the center of each
dorsal scale row. If such a pattern is the result of ontogenetic
change in these northern Haitian D. stenurus, then the dorsal
patterns of the very large specimens is characteristic only of
them and not of merely large-sized D. stenurus.

This pattern achieves some importance, since there are no
meristic scale characters which separate the large species from
its smaller relatives. This, however, is not surprising; examina-
tion of the data from the costatus complex (Schwartz, 1964)
shows that various scale counts have little or no significance
in separating the species stenurus, costatus and curtissi from
each other. For instance, ventrals scales between the mental
and the vent range from 81 to 105 in stenurus, 77 to 100 in
costatus, and 80 to 102 in curtissi; similarly, scales around the
body at midbody are 37 to 45 in stenurus, 35 to 44 in costatus,
and 33 to 42 in curtissi. More important is keeling of the

780 Proceedings of the Biological Society of Washington

dorsal body and caudal scales, and striation of the ventral
scales. Of the three well-known species, stenurus has keeled
dorsal body and caudal scales and smooth ventrals, whereas
both costatus and curtissi have smooth dorsal body and caudal
scales and striate ventrals. The new northern Haitian species
has very strongly keeled dorsal body and caudal scales, and
striate ventral scales, and thus differs in this combination of
characters from all other members of its complex in Hispani-
ola. Differences between these Haitian specimens and the Ja-
maican D. occiduus will be pointed out below; despite the
fact of large size in both occiduus and the new species, the
former is much the larger and bulkier lizard, and there is no
close resemblance or relationship between the two species.

Aside from examining specimens of other Hispaniolan spe-
cies in the ASFS, I have borrowed two specimens of D. oc-
ciduus (UMMZ 53249, 53251) from the University of Michigan,
and one from the Museum of Comparative Zoology (MCZ
74090); I acknowledge the cooperation of Charles F. Walker
and Ernest E. Williams for the loans of such valuable material.
C. Rhea Warren was instrumental in the collection of and my
receipt of one individual of the new form. The second Warren
lizard had been deposited in the living herpetological collec-
tion of the New York Zoological Society, under the curatorship
of F. Wayne King. Dr. King very generously allowed me to
preserve this remarkable lizard, and it has been placed in
the collection of the American Museum of Natural History
(AMNH). I have borrowed the Abbott specimen in the
United States National Museum (USNM) through the co-
operation of James A. Peters and George R. Zug. Mr. War-
ren’s travels in Haiti were greatly facilitated by M. Ramah
Théodore, Directeur Général Adjoint de Office Nationale du
Tourisme et de Propagande. I am especially grateful to Dr.
King for allowing me to preserve a lizard which might other-
wise have afforded an unusual display. In honor of Mr.
Warren, who was instrumental in securing two of the three
known specimens, I name the new species

Diploglossus warreni new species

Holotype: AMNH 103215, an adult female, from Palmiste, Ile de la

A new Diploglossus 781

Tortue, Département du Nord Ouest, Haiti, taken 27 January 1968 by
natives for C. R. Warren. Original number ASFS V15082.

Paratypes: USNM 59435, an adult male, from Riviere des Barres, Dépt.
du Nord Ouest, Haiti, 21 February 1917, W. L. Abbott; ASFS V15071,
same locality as holotype, November 1968, natives for C. R. Warren.

Definition: A species of Diploglossus of the Hispaniolan costatus
complex which differs from other members of that complex (costatus,
stenurus, curtisst) in much larger size (male warreni to 230 mm snout—vent
length; female warreni to 227 mm snout—vent length; largest male and
female of three other species those of stenurus with maximally sized male
172 mm, female 143 mm), low number of ventral scales between mental
and vent (84 to 92 in warreni, 77 to 106 in other members of the complex
combined), dorsal trunk scales striate and with a strong median keel,
dorsal caudal scales likewise striate and strongly keeled, ventral scales
finely striate; dorsal pattern consisting of a series of medium dark
grayish chevrons, more or less confluent medially (especially anteriorly )
and extending the full length of the unregenerated portion of the tail, on
a pale tan to yellow-tan ground, sides very pale gray, flecked with dark
brown and with remnants of vertical bars corresponding to lateral ends of
dorsal chevrons, postocular dark mask obsolete and only very faintly
indicated, iris brown, and venter immaculate cream to very pale orange-
tan.

Distribution: Known only from Ile de la Tortue and from the ad-
jacent Haitian mainland in the Département du Nord Ouest.

Description of holotype: An adult female with snout—vent length of
218 mm, tail (regenerated for distal half) 111 mm; scales between mental
and vent 92, scales around body at midbody 37, 11 chin shields, angular
subocular between supralabials 6 and 7 on both sides; fourth toe
lamellae 15, arm length 32 mm, head width 27.0 mm, head length 34.1
mm. Color in life dorsally yellow-tan (PI. 14 K 6; all color designations
from Maerz and Paul, 1950) with a series of about 16 widely opened
darker brown (Pl. 15 C 9) chevrons from the neck to the groin, the
anterior neck with a vague pair of broad paramedian lines, which ex-
tend to just above the forelimb insertion and are concolor with the dorsal
chevrons and obviously a part of that pattern sequence; about 6 chevrons
between the groin and the regenerated portion of the tail; head scales
tan, immaculate; sides pale gray, flecked with brown especially in the
regions where a very vague series of vertical tannish lateral bars corre-
sponds to the position of the lateral ends of the dorsal chevrons; both
fore- and hindlimbs yellowish tan with a brown reticulum which
isolates islands of the paler ground color; postocular mask obsolete, rep-
resented only by a vague dusky area from the eye onto the temple; upper
Jabials and chin pale gray, with some brownish suture-following pigment
on the chin; venter pale orange-tan (Pl. 12 C 5), immaculate except for
some dark gray blotches on the free edge of the anal flap; subcaudal
scales concolor with those of venter; iris brown.

Variation: USNM 59435, an adult male, has the following measure-

782 Proceedings of the Biological Society of Washington

ments (in millimeters) and counts: snout—vent length 230, tail about
70, distal third regenerated; scales between mental and vent 84, scales
around body at midbody 37, 9 chin shields, angular subocular between
supralabials 6 and 7 on both sides; fourth toe lamellae 18, arm length 48;
head too badly damaged for measurements but conspicuously swollen
and enlarged in masseteric region; specimen badly discolored but dorsal
crossbars or chevrons still barely visible and uncountable; venter com-
pletely immaculate. ASFS V15071, an adult female, has the following
measurements and counts: snout—vent length 227, tail about 175, distal
third regenerated; scales between mental and vent 88, scales around
body at midbody 37, 11 chin shields, angular subocular between
supralabials 7 and & on left side, between 6 and 7 on right side; fourth
toe lamellae 15, arm length 50, head width 32.4, head length 40.7.
Color in life medium brown above with a series of grayish brown
chevronate elements, at times broken middorsally to give a middorsal
reticulum, extending from nape onto unregenerated portion of the tail,
about 18 chevrons between the nape and the groin, about 14 chevrons
between the groin and the regenerated portion of the tail; head scales
medium brown margined with dark brown along their posterior borders;
sides gray, irregularly flecked with brown; limbs slightly darker gray,
the forelimbs with an irregular brownish reticulum, the hindlimbs with
each scale outlined with dark brown; postocular mask obsolete, repre-
sented by an area of irregularly darkened scales in the temporal region;
upper labials grayish, margined with dark brown; venter cream to white,
immaculate except for very pale gray sutures on the sublabials and
throat scales, and irregular black blotches on the free edge of the anal
flap; subcaudal scales with a pattern of dark brown lines radiating
from the base of each scale to give a characteristic dark fan-shaped
pattern.

Comparisons: D, warreni requires comparison only with D. stenurus,
apparently its closest relative on Hispaniola, and with D. occiduus in
Jamaica, the only Antillean species which is as large as or larger than
D. warreni. Comparison with occiduus! is the more easily made (but I

1 Dr. Walker advised me that the smaller of the two UMMZ D. occiduus (UMMZ
53251) has been considered a representative of Diploglossus hewardi. The lizard
may well be hewardi, but I consider it occiduus. Comparison of this specimen with a
series of 22 hewardi suggests the following differences between them and the
problematical UMMZ specimen. Half-jaw counts of position of the angular sub-
ocular in hewardi are regularly 6/7 or 7/8, with the latter category having the
higher incidence (30). The UMMZ lizard has this scale between supralabials 8
and 9 unilaterally, apparently an occiduus character. The two giant occidwus have
striate but unkeeled dorsal scales, weakly striate caudals, and smooth ventrals,
whereas the series of hewardi has striate and strongly keeled dorsals, caudals
striate or striate to very weakly keeled, and ventrals smooth to weakly striate. The
absence of keels on the dorsals of UMMZ 53251 suggests that this specimen is an
occiduus. There seem to be no differences in the head scutellation (other than the
position of the angular subocular) useful in differentiating these two taxa; the
ventral count of 121 in the problematical specimen is slightly higher than the 107

A new Diploglossus 783

doubt that these two species are at all closely related); ventral scales
in occiduus vary between 107 and 121, midbody scales are 49 and 50,
the angular subocular lies between supralabials 8 and 9 (unilaterally in
UMMZ 53251 and MCZ 74090), fourth toe lamellae vary between 16
and 23, the dorsal body and caudal scales are striate but not keeled,
and the ventrals are smooth. UMMZ 53249, an adult male, has a
snout—vent length of 305, head length 66.8, head width 55.5, and arm
length of 69. Similar measurements on MCZ 74090, a female, are 256,
52.9, 40.5 and 57; both are very much larger and bulkier lizards than
D. warreni. All specimens of D. occiduus are presently bleached, so
that no data are available from them on color or pattern. Boulenger
(1885:290) stated that D. occiduus was “Brownish above, with dark
brown spots or cross bands.” Structurally, occiduus differs most strik-
ingly from warreni in that the former has striate dorsal body and
caudal scales and smooth ventrals, whereas warreni has strongly keeled
dorsal body and caudal scales and finely striate ventrals. The higher
number of ventral scales and midbody scales in occiduus likewise sep-
arates the two species; the position of the angular subocular between
supralabials 8 and 9 in occiduus differs from its position between 6 and 7
in warreni.

D. warreni differs from D. stenurus in several features. First, the
strongly keeled dorsal body scales and the very strongly carinate caudal
scales differ from the keeled condition in stenurus. Although these
scales are keeled in stenurus, even in large adult individuals of the latter
species the keels are much lower. This is most especially shown by the
caudals in warreni; these scales have such high keels that the tail appears
angulate on gross inspection. The ventral scales of warreni are finely
striate, whereas these scales are smooth in stenurus. Secondly, the dorsal
pattern, although reminiscent of that of stenurus, is distinctive. D. sten-
urus, D. costatus and D. warreni have a community of dorsal pattern
elements; the pattern is composed basically of chevrons or (when these
are widely opened) crossbars which extend from the nape to at least
the base of the tail. In costatus the chevrons are fine and narrow,
giving a herringbone pattern, whereas in stenurus the chevrons are frag-
mented and are made up of more or less isolated dark squares or
rectangles arranged in a chevronate pattern (see Schwartz, 1964:15,
figs. 1-4), which, however, lacks the clarity and diagrammatic distinct-
ness of the costatus pattern. Both these species likewise have, in
most subspecies, a pair of anterior paramedian nuchal lines, much
better delineated in D. stenurus (especially in D. s. alloeides Schwartz )

and 118 counts for the undisputed occiduus and within the range of that count
in hewardi (113-135), but then very little is known about the variation of this
character in occiduus. Midbody counts in the two large occiduus are 49 and 49-59
in hewardi; the smaller occiduus has a count of 50. I do not consider, nor did
Cousens (loc. cit.), that occiduus and hewardi are conspecific. But immature
occiduus may be difficult to separate from hewardi except for modal differences,
and the same may well be true of juvenile warreni and stenurus.

784 Proceedings of the Biological Society of Washington

than in D. costatus. The dorsal pattern of D. warreni likewise is basically
chevronate, but the chevrons, since they are widely opened, are almost
crossbars. Additionally, these bars are broad and entire (not frag-
mented as in stenurus) and may be joined medially at their apices in an
irregular fashion. There is no indication of paramedian nuchal lines,
although the holotype has dark paramedian blotches on the neck. Thus,
although the species warreni, stenurus and costatus are all basically
chevronate, the degree and quality of the dorsal markings varies with
the species. Thirdly, D. warreni is by far the largest member of the
costatus complex in Hispaniola. Maximally sized stenurus (males first,
females second in each case) are 172 and 143, maximally sized costatus
measure 127 and 116, and maximally sized curtissi 86 and 82. The
male warreni has a snout—vent length of 230, the larger female 227; in
these measurements warreni ranks second only to occiduus in the An-
tilles.

Scale count differences between warreni and stenurus are difficult
to assess, since there are counts available on only three warreni in
contrast to those from several hundred stenurus. However, the warreni
counts of ventral and midbody scales fall toward the lower extremes of
these counts in D. stenurus, and in the case of midbody scales, the
counts of warreni (33-37) lie just below the counts for stenurus (37-45).
Overlap of counts is of little significance in this group of galliwasps,
since even such strikingly different species of stenurus and curtissi have
almost identical extremes in ventral and midbody counts. It would be
extremely pleasant if warreni differed quantitatively from stenurus in
some meristic character, but such is not the case, at least in those counts
which I have heretofore employed in differentiating members of this
Hispaniolan complex.

One other characteristic separates warreni from stenurus. In the
former, the subcaudal scales are very large and almost fanlike in aspect;
this resemblance is further enhanced by the dark brown lines of pigment
which may radiate from the base of each scale. Such radiations do not
occur in stenurus and the ventral caudal scales are relatively (as well
as actually) much smaller. Thus the subcaudals in warreni are much
larger than are those in stenurus. Finally, the distinctly angulate appear-
ance of the tail, due to the high median keels on the dorsal caudal
scales, does not occur in stenurus; in the latter species, although the
dorsal caudals are keeled, the keels are not so high and do not impart
an angulate or longitudinally keeled aspect to the upper side of the tail.

Remarks: Despite the fact that I regard D. warreni as a species
distinct from D. stenurus, nevertheless, the very fact that I previously
considered the USNM paratype of D. warreni as a stenurus suggests
the similarity between these two species. The age and condition of the
USNM specimen, as pointed out above, prevented my assessment of its
characteristics. Still, D. stenurus is unknown from Ile de Ja Tortue and
from the northern Haitian coast in the immediate region of St. Louis
du Nord-Riviére des Barres. Thus, there is a possibility that the speci-

A new Diploglossus 785

mens I consider D. warreni are either 1) merely very large D. stenurus
or 2) represent a local very large subspecies of D. stenurus. Neither
of these possibilities can be absolutely refuted. But the facts that large
stenurus from adjacent areas show no approach to the dorsal pattern of
warreni and that, of hundreds of stenurus studied from throughout
Hispaniola, none begins to approach the very large size of warreni
suggests that these lizards are related to, but not conspecific with, D.
stenurus. The structural characteristics of D. warreni, in comparison
with those of D. stenurus, of course also militate against the conspec-
ificity of the two taxa. The compact known geographic distribution of
D. warreni also suggests that this population is distinctive from D.
stenurus.

There is another inductive line of reasoning for specific status of
D. warreni. Etheridge (1965:99-100), while discussing the fossil lizards
from a cave at Cerro de San Francisco near Banica in San Rafael Prov-
ince, Reptblica Dominicana, distinguished between two size-classes of
fossil Diploglossus found in this cave: one group with a presumed
snout—vent length of 120 to 130 mm (costatus) and another group with
a presumed snout—vent length of about 210 to 250 mm (“stenurus’).
Dr. Etheridge made the latter assignment at the time because of my
as yet unpublished data on the maximum size of D. stenurus; this
maximum size (230 mm) was of course based upon the USNM para-
type of D. warreni. The presence in these deposits of the two size
groups suggests that there were indeed two species present, one of
which (warreni) is much the larger. D. warreni is not known today
from the Banica area.

The presence of apparent warreni fossils from the interior of the
Republica Dominicana, and the known occurrence today of this species
only along the northern Haitian littoral and on Ile de la Tortue suggest
that the species was once more widely distributed, and that it persists
today mainly along the northern Haitian coast and on Tortue. The
absence of D. stenurus on Tortue is especially puzzling, but considering
the apparent rarity of any species of Diploglossus on that island, it is
possible that stenurus occurs there. Another possibility is that large
stenurus is incapable of competing with even larger warreni, and that
stenurus has not been able to establish itself on Tortue where warreni
is a long-established species. If the Banica fossils are correctly inter-
preted as warreni, the previously widespread distribution of that species
and its present apparently circumscribed (and partially insular) range
seems clearly to indicate that warreni is a relict species, persisting in
smail numbers in northern Haiti and on Ile de la Tortue.

Mr. Warren advises me that the vicinity of Palmiste is mesic, and that
the region immediately about the Riviére des Barres is likewise mesic,
sufficiently so to allow the growing of bananas and coffee very near
the coast. Farther to the west at Port-de-Paix, the countryside is
scrub-grown and arid. Of the lizards reported by Etheridge from the
pre-Columbian strata at Cerro de San Francisco, none is a confirmed

786 Proceedings of the Biological Society of Washington

inhabitant of xeric areas (although A. chrysolaema is a xerophile in
some parts of its range); several species (Anolis ricordi, Anolis cybotes,
Anolis chlorocyaneus, Leiocephalus personatus, Ameiva taeniura, Diplo-
glossus costatus) are inhabitants of cool, shaded, and relatively moist
areas. Considering the habitat preference (so far as it can be known
from the few specimens available) of living D. warreni, it seems possible
that the region about Cerro de San Francisco was formerly mesic and
forested.

In summary, it seems probable that D. warreni was once more widely
distributed on Hispaniola, but that (because of stringent ecological
requirements?) it has disappeared from much of its former range and
persists only on the mesic northern Haitian coast and on Ile de la
Tortue. Perhaps the presence of D. warreni on Tortue has prevented
the success there of its largest congeneric relative, D. stenurus.

Underwood (1959:11) suggested that since the Jamaican micro-
blepharis was related to Cuban-Puerto Rican delasagra—pleii it was most
likely that this group of species, on the basis of geographical distribution
of its known members, must have a Hispaniolan representative. I
agree, but D. warreni is not that as yet unknown member. Rather, the
relationships of warreni are clearly with the stenurus—costatus—curtissi
series on Hispaniola, warreni representing the culmination of size in this
complex. Similarly, occiduus in Jamaica seems to represent the cul-
mination of a Jamaican series (crusculus-hewardi-barbouri) with crus-
culus the smallest member in the series. In both cases the series pro-
gresses from small short-limbed species to very large long-limbed
species. As in the costatus complex, the basic Jamaican dorsal pattern
is a series of fine and narrow, closely appressed chevrons, giving a
herringbone pattern. I am less sure of this sequential series in Jamaica
than in the costatus series in Hispaniola; for one reason, Underwood
(op. cit.:13, table) indicated that occiduus is the only member with
many scale organs on the dorsal scales whereas all other Jamaican galli-
wasps have the scale organs present on the dorsal scales but apparently
they are not so numerous as in occiduus. If many scale organs is a more
primitive condition than few-to-no scale organs, then on the basis of
this character, occiduus would seem to be the primitive member of
the Jamaican series, and the remainder of the Jamaican species are
derived forms—a situation which, on the basis of size, seems unlikely.
Thus, the series of Jamaican galliwasps does not demonstrate the
clearly sequential evolutionary series that the Hispaniolan costatus
group members show.

LITERATURE CITED

BouLencer, G. A. 1885. Catalogue of the lizards in the British
Museum (Natural History). London, 2: 1-498.

Cocuran, Doris M. 1941. The herpetology of Hispaniola. Bull. U. S.
Nat. Mus., 177: i-vii, 1-398, 12 pls., 120 figs.

Cousrens, PENNy N. 1956. Notes on the Jamaican and Cayman Island

A new Diploglossus 787

lizards of the genus Celestus. Breviora, Mus. Comp. Zool.,
56: 1-6.

Eruermwce, RicwArp E. 1965. Fossil lizards from the Dominican
Republic. Quart. Jour. Florida Acad. Sci., 28(1): 83-105,
3 figs.

GraNtT, CHAPMAN. 1940. The herpetology of Jamaica, II. The Rep-
tiles, in Lynn and Grant. The Herpetology of Jamaica. Bull.
Inst. Jamaica, 1: 61-147, 3 figs.

Maerz, A., AND M. RuHeEA Paut. 1950. A dictionary of color. New
York, McGraw-Hill Book Co., pp. i-vii, 1-23, 137-208,
56 pls.

Scuwartz, ALBERT. 1964. Diploglossus costatus Cope (Sauria, An-
guidae) and its relatives in Hispaniola. Reading Public
Mus. and Art Gallery, Sci. Publ. 13: 1-57, 20 figs.

UNDERWOOD, GARTH. 1959. A new Jamaican galliwasp (Sauria, An-
guidae). Breviora, Mus. Comp. Zool., 102: 1-13, 4 figs.

788 Proceedings of the Biological Society of Washington

I7Y.067F

Vol. 82, No. 61, pp. 789-798 5 February 1970

PROCEEDINGS
OF THE

BIOLOGICAL SOCIETY OF WASHINGTON

A NEW IDOTEID ISOPOD, IDOTEA (PENTIDOTEA)
KIRCHANSKII, FROM CENTRAL CALIFORNIA
(CRUSTACEA )

By Mitron A. MILLER AND WELTON L. LEE
Department of Zoology, Davis Campus, University of
California and Hopkins Marine Station, Stanford University

In his monograph on the idoteid isopods of northern
California, Menzies (1950) reduces the genus Pentidotea
Richardson, 1905, to the rank of a subgenus of Idotea’ Fab-
ricius, 1798, and gives an account of the six species then known.
The present paper describes a seventh species in this subgenus.
It is the third species of Idotea (Pentidotea) associated with
surf grass (Phyllospadix scouleri), the other two being I.
(P.) montereyensis (Maloney) Menzies, 1950 and I. (P.)
aculeata (Stafford) Menzies, 1950. The new idoteid was first
discovered during the course of ecological investigations on
Idotea montereyensis (Lee, 1967) living alongside it on Phyl-
lospadix, but unlike the latter isopod, Idotea kirchanskii did
not appear to occur elsewhere intertidally.

Genus Idotea Fabricius, 1798

Menzies (1950) gives an emended diagnosis of the genus as follows:
“GENERIC DIAGNOSIS: Flagellum of second antennae multiarticu-
late. Maxillipeds with a palp composed of four or five articles. Epimera
of all the segments (somites) of thorax (peraeon), with the exception
of the first, distinctly separated from the somites. Abdomen (pleon)
composed of three segments, with a suture line on either side at the base
of the terminal segment, indicating perhaps another partly coalesced».
segment. Includes the subgenera Idothea and Pentidotea.” i =

i
1 Many authors, including Richardson (1905) and Menzies (1950), spell the
generic name “Jdothea” but Fabricius’ original spelling was ‘“‘Idotea.” a

61—Proc. Biot. Soc. WasuH., Vou. 82, 1970 (789 ) \S.

790 Proceedings of the Biological Society of Washington

Subgenus Pentidotea (Richardson) Menzies, 1950

Maxillipedal palp with 5 articles rather than 4 as in the subgenus
Idotea.

Idotea (Pentidotea) kirchanskii new species
Figures 1 and 2

Diagnosis: Body narrow, linear, compact. Color bright green with
tips of appendages often red. Frontal process of head broadly triangulate
with bluntly rounded apex, shorter than frontal lamina 1 of clypeus.
Frontal lamina 1 prominent, broadly triangulate, wider and longer than
frontal process and concealing frontal lamina 2 in dorsal view. Antenna
2 short, rarely extending past posterior margin of pereonite 2, with
flagellum comprising 7-12 articles. Eyes round. Maxilliped with 1
coupling hook, rarely 2. Epimera distinctly separated on pereonites 2-7,
but usually visible dorsally only on segments 5—7. Epimera of pereonite 7
triangular with acute posterior angles. Pleotelson with medial posterior
margin convex, lacking apical tooth. Found on Phyllospadix.

Description: The following characters may be noted in addition to
those given above.

Holotype male (21 mm x 3.2 mm), allotype ovigerous female (15
mm X 2.8mm). Paratypes: 5 males, 1 nonovigerous female, 8 ovigerous
females. In type series, length/width ratios in six males range between
5.2 and 7.1, in the nonovigerous female, the ratio is 5.6; in nine
ovigerous females, the range is 4.3-5.7 due to expansion in width at
pereonites 1-3.

Head (Fig. 1, a—b) with supra-antennal line slightly concave, but
median third of it often becomes convex. Anterolateral cephalic margin
slightly flared, rounded. Eyes round (lateral view). Lateral margins
slightly indented anterior to eyes. Frontal lamina 2 of clypeus broad and
almost semicircular in frontal view.

Antenna 1 (Fig. 1, c) composed of 4 short articles, extending only to
about two-thirds the length of third peduncular article of antenna 2. First
article of antenna 1 stout, about as wide as long, and twice as broad as
those following; articles 2-4 subequal in length; article 4 somewhat
clavate and provided with stout apical and subapical setae (sensory?).
Antenna 2 (Fig. 1, d) rarely extends past posterior margin of second
pereonite. Peduncle of antenna 2 composed of 5 articles, the first
small and barely visible dorsally; articles 2 and 3 larger and subequal;
articles 4 and 5 narrower and each slightly longer than third article.
Flagellum of antenna 2 consists of 7-12 articles (but juveniles may have

>

Fic. 1. Idotea (Pentidotea) kirchanskii new species. a, ¢ from

Dillon Beach, dorsal view. b, head with left antennae and postmandib-

ular mouthparts removed, side view. c, first antenna. d, second antenna.

e, first pereopod. f, seventh pereopod. g, left mandible. h, first maxilla.
i, second maxilla. 7, maxilliped.

A new California isopod

791

792 Proceedings of the Biological Society of Washington

as few as 3 joints)—the first longest and often showing partial articu-
lation; the last minute and semicircular in outline.

Mouthparts normal for genus (Fig. 1, gj). Mandibles with heavily
sclerotized incisive process and truncate molar process bearing a brush
of setae. First maxilla has inner lamina with 3 long plumose setae and
one short seta at apex, and outer lamina with stout apical spines, the
inner slightly ctenate. Second maxilla trilobate, inner lobe fringed with
long plumose setae, outer lobes with ctenate setae. Maxilliped has
palp of 5 articles, and endite with 1 coupling hook (rarely 2).

Lateral thoracic margins subparallel, not sharply incised between seg-
ments. Posterior margin of first pereonite decidedly concave, that of
second less so, that of third more or less straight, those of following
segments medially convex.

Epimera of pereonites 2-4 do not extend the length of their segments,
whereas those of segments 5—7 do. Epimera of pereonites 2 and 3 never
visible from above, those of pereonite 4 sometimes visible dorsally, those
on remaining three pereonites at least partially visible dorsally. Postero-
lateral edges of fifth and sixth segments somewhat rounded, edges of
seventh form an acute angle. Sternal parts of coxal rings not fused
medially leaving prominent medial groove. Coxal sockets deep.

Pereopod 1 (Fig. 1, e) bears row of stout conical setae along entire
palmar margin of propodus; each seta complex with many fine filelike
ridges facing palm and a filiform extension of its axis beyond bifid apex.
Also one large complex seta and several smaller simple setae on distal
edge of carpus. Dactylus with long curved unguis and accessory claw at
base. Pereopod 7 (Fig. 1, f) with keel on external margin of basis, and a
few stout, coarsely pectinate, spiniform setae at base of palmar margin of
propodus and a few at distal margin of carpus.

Posterior margin of telson slightly convex, evenly rounded, lacking
posterolateral angles, sometimes slightly produced in broad median lobe,
but without apical tooth.

Penes (Fig. 2, a) double, short, flattened, bluntly tapered processes
attached to sternite of seventh pereonite. Pleopod 1 (Fig. 2, b) with
quadrate base bearing 8 stout coupling spines on medial distal border and
two laminar rami each fringed with long plumose setae. Pleopod 2 of
male (Fig. 2, c) bears long, slender appendix masculina extending
along entire length of medial edge of endopod and beyond its distal
edge, with rounded apex armed subapically with rows of spiny scales;
both rami fringed with plumose setae. Posterior 3 pairs of pleopods also
biramous, but show progressive reduction in marginal setation. Pleopod
3 (Fig. 2, e) has setae only on exopod and these are sparse compared
to those of first and second pleopod, with plumose types limited to
distal and distolateral margins. Pleopods 4 and 5 (Fig. 2, f-g) similar,
with rami seemingly bare, but under magnification exopods show a
sparse fringe of short, spinelike setae. Branches of pleopods 1 and 2
uniarticulate, but exopods of last 3 pairs show lateral and medial partial
sutures connected by fine transverse lines making them biarticulate.

A new California isopod 793

LOmm

f g

Fic. 2. Idotea (Pentidotea) kirchanskii new species, ¢ pleonal
appendages. a, penes. b, first pleopod. c, second pleopod. d, tip of
appendix masculina. e, third pleopod. f, fourth pleopod. g, fifth pleopod
(exopod somewhat flattened). h, uropod.

Uropod (Fig. 2, h) uniramous with 3 plumose setae at outer distal
angle of basal joint.

Localities: CALIFORNIA: Monterey County, Pebble Beach, 17 Mile
Drive at Seal Rock, intertidal on Phyllospadix (6 ¢ 6,10 92 [9 ovig-
erous]), 14 May 1968, W. L. Lee (type-locality ); Marin County, Dillon
Beach, Second Sled Road, intertidal on Phyllospadix (10 6&6, 7 @Q
[2 ovigerous]), 18 June 1962, W. L. Lee. Additionally, this isopod has

794 Proceedings of the Biological Society of Washington

TABLE 1. Similarities between Idotea kirchanskii and I. aculeata.

1. Body linear, sides subparallel.

2. Frontal process widely angulate and shorter than frontal lamina 1,
with blunt, evenly rounded apex. In ovigerous females of I. aculeata,
Menzies (op. cit.) notes that the apex is often somewhat concave, a
variation not found in I. kirchanskii.

3. Frontal lamina 1 prominent, broadly triangulate, wider than frontal

process and extending forward beyond it in dorsal view.

Eyes round.

Single coupling hook on maxilliped.

First pleonite with wide lateral borders.

Posterolateral margin of epimeron of seventh pereonite acute.

Posterolateral angles of pleotelson rounded.

il ad

been collected intertidally on many occasions at Dillon Beach, Marin Co.;
and in Monterey Co., at Pacific Grove and Pebble Beach and in various
localities between Carmel and Rocky Point. It has only been found on
Phyllospadix scouleri and invariably it is the same green color as that
plant.

Disposition of material: Types are deposited in the Smithsonian Insti-
tution. Holotype—USNM 125205, allotype—USNM 125206, paratypes
—USNM 125207. Other material has been placed in the collections of
the authors and the California Academy of Sciences, Golden Gate Park,
San Francisco.

Etymology: The species is named in honor of Mr. James Kirchanski
in recognition of his contribution to education in central California.

Distribution: The known geographical range of Idotea kirchanskii
extends south from Dillon Beach, Marin County to Rocky Point, Mon-
terey County, California. Its distribution within this range is imper-
fectly known, but it appears to be more abundant in the southern portion.
Relatively few specimens were found at Dillon Beach where I. monte-
reyensis is the dominant species on surf grass. At Rocky Point, however,
I. kirchanskii is abundant and almost completely replaces I. monterey-
ensis.

The range of Idotea kirchanskii overlaps the northern end of the
distribution of its closest relative, Idotea aculeata, which is found from
Dillon Beach, Marin County, south to La Jolla, San Diego County, Cali-
fornia (Menzies, 1950). George and Stromberg (1968), however, report
I. aculeata from San Juan Archipelago, Washington, a locality far north
of its previously known range. I. aculeata, according to Menzies, is a
major component of the isopod fauna south of Point Conception—a
major breakpoint at about 341° N latitude which separates the temperate
marine biota of northern California from the transitional warm tem-
perate biota of southern California. It is possible that some reports of
I. aculeata north of Point Conception may be I. kirchanskii as the two
species are quite similar (see Table 1).

A new California isopod 795

TaBLe 2. Differences between Idotea kirchanskii and I. aculeata.

I. kirchanskii

1. Body
2. Color
3. Head

4, Antennae 2

5. Pereonite 3

6. Epimera

7. Pleonites

8. Telson

I. aculeata

Compact, pereonites
without lateral inci-
sions between them.

Always green.

Eyes bulge at
lateral margins.
Frontal process
not concave.

Short, barely reach-
ing posterior margin
of pereonite 2.

Posterior margin
more or less
straight.

Dorsally visible only
on pereonites 5-7.

Lateral margins
curved inward
anteriorly.

Posterior margin
without apical
tooth.

Not compact, lateral tho-
racic margins incised,
especially between
pereonites 1-4,

Mostly pink or red.

Lateral margins not
bulged. Frontal process
in ovigerous females
often concave.

Longer, reaching almost
to posterior margin of
pereonite 4.

Posterior margin concave.

Dorsally visible on
pereonites 2—7.

Lateral margins entirely
straight.

Posterior margin with
bluntly pointed median
tooth.

Relationships: The fact that Idotea kirchanskii keys to I. aculeata in

Menzies’ (1950) monograph indicates a close morphological relationship
between these two species. They differ significantly, however, in other
respects. The similarities between the two species are shown in Table 1,
the differences in Table 2.

A few specimens which have characters intermediate between Idotea
kirchanskii and I. montereyensis have been found. I. montereyensis is
also found on the same plant as the new species which suggests the
possibility of some hybridization between them.

The separation of I. aculeata from other pentidoteans in Menzies’
(1950) key is based on characteristics common to both I. aculeata and
I. kirchanskii; however, with few modifications the key can be
altered to conveniently include I. kirchanskii. The emended key is
presented below:

796 Proceedings of the Biological Society of Washington

KEY TO THE NORTHERN CALIFORNIA SPECIES OF
THE SUBGENUS PENTIDOTEA

1. Apex of frontal process entire. Maxilliped with one coupling
hook. Eyes not markedly transversely elongate, length along

body axis one-half or greater than one-half the width ____ 2
— Apex of frontal process with a median notch — 6
2. Frontal process blunt or widely angulate, not extending beyond

frontal lamina 1. Frontal lamina 1 triangulate in dorsal view. 3
—A narow, pointed frontal process exceeds considerably the for-

ward extent of a semicircular frontal lamina ] — 5

3. Posterolateral margin of epimeral plate of seventh peraeon
somite evenly convex, not acute. Eyes somewhat pyriform
Sot ete SON CN Oe Coe ar ee ee I. (P.) schmitti Menzies

— Posterolateral margin of epimeral plate of seventh peraeon somite
acute. Eyes reniform or oval _..- 4

4, First pleon somite with acute lateral borders. Eyes reniform
ac RR ei atte Roe I. (P.) wosnesenskii (Brandt) Menzies

— First pleon somite with wide lateral borders. Eyes circular _. 7

5. Telson posterior margin deeply concave, posterolateral angles
acute, each angle with a small but noticeable dorsal carina.
Specimens usually found on eel grass (Zostera sp.) ------
posi sstchpte pean nestdacicncchieeenahoatamagemabeimnne oe I. (P.) resecata (Stimpson)

— Telson posterior margin usually convex, with a small but distinct
median tooth; when concave then only slightly so and lack-
ing acute posterolateral angles and lacking any dorsal carina
above each angle. Specimens usually found on surf grass
(Phyllospadix sp.) —— I. (P.) montereyensis (Maloney )

6. Maxilliped with two coupling hooks. Eyes transversely elongate
Fee ee eee oe NI MeN ea I. (P.) stenops (Benedict)

— Maxilliped with one coupling hook. Eyes oval —----------.--
cpadiverbianntedl Some adult specimens of I. (P.) aculeata (Stafford)

7. Segments distinctly separated laterally, lateral margins of ceph-
alon entire, second antennae reaching almost to posterior
margin of fourth thoracic segment, epimera noticeably visible
dorsally on segments two through seven _...----
One ee MOONEE A Ee ERE eee AME ree SO I. (P.) aculeata (Stafford)

— Segments not sharply separated laterally, lateral margin of ceph-
alon indented anterior to the eyes, second antennae short,
reaching almost to posterior margin of second thoracic seg-
ment, epimera noticeably visible dorsally only on segments
five, six and seven ____...- I. (P.) kirchanskii Miller and Lee

Ecology: Little is known of the ecology of this species. It appears,
however, to be restricted to one particular habitat, the green intertidal
flowering plant, Phyllospadix scouleri. It is remarkably adapted for
clinging to the wave-swept blades of this surf grass and accordingly is
found on it even in rough water. Analysis of gut contents suggests that

A new California isopod 797

the animal feeds on both Phyllospadix and a wide variety of epiphytes
found on the long narrow blades of the plant.

Initial investigations suggest that Idotea kirchanskii does not seem to
change color in response to a change in the color of its substrate as does
I. montereyensis (Lee, 1966a, 1966c), although superficial examination
has revealed a pigmentary system similar to that found in the latter
species (Lee, 1966a, 1966b).

LITERATURE CITED

Grorce, R. Y., AND J-O. STROMBERG. 1968. Some new species and
new records of marine isopods from San Juan Archipelago,
Washington, U.S.A. Crustaceana, 14(3): 225-254.

Ler, W. L. 1966a. Pigmentation of the marine isopod Idothea mon-
tereyensis (Maloney). Comp. Biochem. Physiol., 18: 17-36.
1966b. Pigmentation of the marine isopod Idothea granu-
losa (Rathke). Comp. Biochem. Physiol., 19: 13-29.
1966c. Color change and the ecology of the marine isopod
Idothea (Pentidotea) montereyensis Maloney, 1933. Ecol-
ogy, 47(6): 930-941.

Menzies, R. J. 1950. The taxonomy, ecology, and distribution of
northern California isopods of the genus Idothea with the
description of a new species. Wasmann Journ. Biol., 8(2):
155-195.

RicHarpson, H. 1905. A monograph on the isopods of North Amer-
ica. Bull. U. S. Nat. Mus., 54: liii + 727 pp., 740 figs. in
text.

798 Proceedings of the Biological Society of Washington

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No. 62, pp. 799-824 5 February 1970

PROCEEDINGS
OF THE

BIOLOGICAL SOCIETY OF WASHINGTON

PELAGIC OSTRACODS (MYODOCOPA HALOCYPRI-
DIDAE) FROM THE NORTH ATLANTIC OFF
BARBADOS!

By GrorciANa B. DEEVEY

Institute of Oceanography, Dalhousie University,
Halifax, N. S., Canada

In connection with a study on primary productivity in the
tropical North Atlantic off Barbados and in the Caribbean
Sea off Jamaica, zooplankton samples were collected and the
related hydrography and nutrient chemistry investigated by
Beers, Steven and Lewis (1965, 1968) in a project carried out
jointly by members of the staffs of the Bermuda Biological
Station, the University of the West Indies in Jamaica, and
the Bellairs Research Institute in St. James, Barbados. The
zooplankton tows collected at the station off Barbados were
stored at the Bermuda Biological Station, and these samples
have been examined for pelagic ostracods. The majority of
the zooplankton hauls were surface tows, collected to obtain
an estimate of the standing crop of zooplankton, but between
September 1963 and May 1964 samples were also collected
at a depth of 400 m. Very few ostracods were found in the
surface samples, but 26 species of Halocyprids were re-
corded from the six 400-m samples obtained in September,
October, November 1963 and January, April and May 1964.

The Barbados station, at 15°12’N, 59°47.5’W, in 450 m‘“of
water, is eight miles west of the Bellairs Research Institute-in
St. James. The zooplankton tows were made with a half-meter
diameter net of No. 8 nylon mesh, equipped with a flow-meter

which recorded the distance towed in meters. The surface ~~

tows, which caught too few ostracods for further considera~ en

1 Contribution No. 480 from the Bermuda Biological Station. fie
62—Proc. Brot. Soc. Wasu., Vou. 82, 1970 (799).

\

800 Proceedings of the Biological Society of Washington

tion, were of 10 minutes duration; the 400-m hauls were towed
for 30 minutes. All the ostracods were removed, counted and
identified from one-fourth of each 400-m sample. In the fol-
lowing listing of the species only those species will be de-
scribed and figured that were not included in my report on the
pelagic ostracods of the Sargasso Sea (Deevey, 1968). The
species found at the station off Barbados, which were not taken
in the Sargasso Sea, include Fellia bicornis (Miller), Concho-
ecia echinata Miiller, C. nasotuberculata Miller, C. parviden-
tata Miller, and Euconchoecia sp.

Ostracods constituted at most 0.4 percent of the total num-
bers of organisms at the surface, but percentages ranged from
0.2 to 6.8 percent in the 400-m samples. The species of ostracods
are listed in Table 1, which also gives the percentages of each
species, based on the total numbers of ostracods. Highest
numbers were found from November to May; few were
present in September and October. Unidentifiable juvenile
ostracods constituted 27-46 percent of the total numbers of
ostracods. The most abundant species were all small, ranging
0.6-1.6 mm in length, and included Archiconchoecia striata,
Conchoecia curta, C. oblonga, C. procera, and C. spinirostris.
Conchoecia acuminata, C. atlantica, C. elegans, C. parthenoda,
C. porrecta and C. rotundata were also relatively numerous
and occurred in every sample. The commonest species at the
Barbados station were also the most abundant in the Sargasso
Sea (Deevey, 1968). One species, C. imbricata, which oc-
curred year-round in the Sargasso Sea, was not taken at
Barbados; other species including C. atlantica, C. acuminata,
C. bispinosa, C. elegans, and C. porrecta, were more numerous
at the Barbados station.

Data for temperature, salinity and nutrient chemistry were
obtained from the surface down to 300 m, but not to 400 m,
during the period studied. At 300 m the temperature range
was 10.51-15.12°C, with a mean of 13.08°C, and the salinity
varied between 35.1 and 35.9%,

The samples were collected under Contract NONR 1135(05)
from the Office of Naval Research. This study was supported
partly by grant GB-2668 and partly by GB-6879, both from the
National Science Foundation.

North Atlantic ostracods 801

SUBORDER HALOCYPRIFORMES Skogsberg 1920
Famity HaLocypripipAE Dana 1852
SUBFAMILY ARCHICONCHOECINAE Poulsen 1969
Genus Archiconchoecia Miiller 1894
Archiconchoecia striata Miiller

Archiconchoecia striata G. W. Miiller, 1894, p. 225, Pl. 6, Figs. 31-46,
Pl. 8, Fig. 34; 1906a, p. 45, Pl. VII, Figs. 13-17; 1912, p. 56.
Archiconchoecia striata, G. B. Deevey, 1968, p. 23, Fig. 4.

This tiny species (males and females are 0.5-—0.6 mm long) was one
of the three most numerous ostracods, present in every sample, and
constituted 2.4-17.3 percent (see Table 1) of the total numbers of
ostracods.

Distribution: 36°N-37°S in the Atlantic, Indian, and Pacific Oceans
and Mediterranean Sea.

SUBFAMILY EUCONCHOECINAE Poulsen 1969)
Genus Euconchoecia Miiller 1890
Euconchoecia chierchiae Miiller

Euconchoecia chierchiae Miller, 1890, p. 277, Pl. XXVIII, Figs. 1-10.
Euconchoecia chierchiae, Skogsberg, 1920, p. 740, Figs. 148-151.
Euconchoecia chierchiae, Deevey, 1968, p. 116, Fig. 62.

Euconchoecia chierchiae, Poulsen, 1969, p. 38, Figs. 12-13.

This species was recorded in September and November 1963 and in
January and April 1964 in small numbers.

Distribution: Atlantic, Pacific, and Indian Oceans between 40°N
and 40°S.

Euconchoécia sp.
(Figure 1)

Three female specimens, differing from E. chierchiae females in ap-
pearance, were taken in the April 1964 sample. The shells (Fig. la—c)
are 1.15, 1.17 and 1.2 mm long by 0.55 mm high, the height of the
shell being around 47 percent of the length, and the greatest height
just behind mid-length. There are no points at the posterodorsal corners
of the shells, and the long point on the left rostrum, characteristic of
E. chierchiae females, is also lacking. In E. chierchiae females from this
station the depth of the shell is less, 38-42 percent of the length, the
greatest height being just anterior to mid-length, there is a long spine at
the posterodorsal corner of the right shell, and the left rostrum is pro-
duced into a sharp point.

The frontal organ is slim, undifferentiated, rounded at the tip, and
reaches almost to the tip of the first antenna (Fig. 1d), which has
distoventrally a cluster of 20-21 sensory filaments, all about the same
length as the principal seta. The frontal organ differs from that of E.
chierchiae in that it is rounded, not pointed nor bifid at the tip; also in
E. chierchiae females the frontal organ reaches to the tip of the

802 Proceedings of the Biological Society of Washington

TABLE 1. Percentages of ostracod species, based on total numbers of

ostracods, from September 1963 to May 1964 in one-quarter of each

400-m sample. X indicates presence of species in one-quarter sample
not counted.

Sept. Oct. Nov. Jan: Apr. May

Archiconchoecia striata 9.78 2.38 17.33 9.00 3.57 10.00
Euconchoecia chierchiae X — 1.93 0.56 1.78 —

Euconchoecia sp. = — — — X —

Halocypris brevirostris — — X 0.56 4.16 1.04
Fellia bicornis Xx — — xX a —

Conchoecia unid. larvae 42.60 46.40 40.00 34.90 27.40 37.50
C. acuminata Xx 2.38 039 028 030 £0.21
C. atlantica Li? 6x 193 253 5.95 0.42
C. bispinosa 0.89 xX X X — 1.04
C. concentrica 2.22, — 0.39 — 0.30 0.21
C. curta 15.58. 15.45 1.93 18.00 12.20 15.20
C. daphnoides X — X 0.56 xX 0.21
C. echinata — Xx — 1.68 0.60 1.67
C. elegans 3.11 1.19 1.16 1.68 0.89 1.46
C. magna L7s 1.19 1.93 2.53 — 1.67
C. nasotuberculata — — — Xx — —

C. oblonga 5.78 5.96 6.55 4.22 10.10 10.00
C. parthenoda L3o ok 694° 225 238 3.96
C. parvidentata X — — — = —

C. porrecta 0.45 2.30 2.70 xX 0.30 0.21
C. procera 10.20 13.10 733 9.57 22.00 10.20
C. rotundata 133 2388 2.31 422 2.68 1.67
C. secernenda 0.89 — 1.93 1.96 0.30 0.21
C. spinifera X 2.38 — 0.56 — —

C. spinirostris 222 4.76 4,24 4.78 5.06 2.08
C. subarcuata Xx — X X — 0.62
Conchoecia sp. xX — — — — 0.42

first antenna. The basal segment of the exopodite of the second antenna
is approximately 44 percent the length of the shaft; the basal segment of
the endopodite is approximately 25 percent shaft length, and bears
dorsodistally two bristles, the shorter of which is slightly more than
half as long as the longer (Fig. le). The distal segment of the endop-
odite has one long seta, which was broken on all three specimens, and
two shorter setae or filaments, the longer one slightly more than twice
as long as the shorter; in E. chierchiae females the longer filament is
less than twice as long as the shorter one. In the morphology of the other
appendages there appear to be no marked differences between these
females and female E. chierchiae. The furca has seven claws, with a

North Atlantic ostracods 803

Fic. 1. Female Euconchoecia sp. a—c, Lateral, ventral and posterior
views of shell. d, Frontal organ and first antenna. e, Endopodite of
second antenna (long seta cut off). Scale at bottom for a—c, at right
for d, e. Scales in mm.

pronounced knob between the first and second claws, and a tiny unpaired
bristle behind the claws. The first several pairs of claws are less
abruptly pointed than in E. chierchiae.

This species is very closely related to E. chierchiae. It is not named at
this time, not only due to lack of more material including males, but
also because a species of Euconchoecia lacking points at the postero-
dorsal comers is presently being described (Tseng, personal communi-
cation).

804 Proceedings of the Biological Society of Washington

SUBFAMILY HALOCYPRINAE Poulsen 1969

Poulsen (1969) separated the genus Halocypris into three genera:
Halocypris, Halocypria, and Fellia. Two species of two of these genera
were found at the station off Barbados.

Genus Halocypris Dana 1852
Halocypris brevirostris (Dana)

Halocypris brevirostris, Skogsberg, 1920, p. 584, Figs. 112-115.
Halocypris brevirostris, Deevey, 1968, p. 19, Fig. 2a—-f, 3c-e.
Halocypris brevirostris, Poulsen, 1969, p. 63.

For synonymy, see Skogsberg.

H. brevirostris was found from November to May, with highest per-
centages in April. This species has a wide range in size of mature
individuals. According to Poulsen (1969) mature females vary in
length from 1.1-2.1 mm, and males from 0.9-1.9 mm. This species
occurs year-round in the Sargasso Sea (Deevey, 1968), where all the
mature specimens were small, females being 1.1-1.3 mm, males 0.95-
1.15 mm long. Mature females from the Barbados station were 1.6 mm,
males 1.4 mm long.

Distribution: 60°N-40°S in the Atlantic, Pacific, and Indian Oceans.
According to Poulsen, it occurs most frequently in the tropics.

Genus Fellia Poulsen 1969:
Fellia bicornis (Miiller)
(Figure 2)

Halocypris bicornis Miiller, 1906a, p. 49, Pl. VIII, Figs. 8-12, 17; 1912,
p. 58.

Halocypris taurina Vavra, 1906, p. 66, Pl. 7, Figs. 128-130, 131, 132a.

Fellia bicornis, Poulsen, 1969, p. 89, Fig. 38.

Poulsen has recently placed in a new genus, Fellia, two species that
were formerly in the genus Halocypris, F. cornuta (Miiller) and F.
bicornis (Miller), with spines and/or rounded processes on the shells.
Both these species occur in all oceans, mainly in tropical regions between
25°N and 25°S;

Ten specimens of F. bicornis were found at the Barbados station in
September, January and February. Three juveniles were 0.6—0.7 mm
long, four were 1.0-1.05 mm long, two were 1.4 mm long, and a
mature female was 1.9 mm long by 1.55 mm high. Lateral, ventral
and posterior views of the female shell, the frontal organ, first antenna,
and the endopodite of the female second antenna are illustrated in
Figure 2.

Distribution: Miiller’s specimens were found between 10°N and 10°S
in the Atlantic and Indian Oceans. Poulsen’s Dana specimens were col-
lected from 45°N-8°S in the Atlantic, in the Indian-Indonesian region
from 23°N-15°S, and in the Pacific from 5°-45°S. Very few specimens

North Atlantic ostracods 805

a
(

aamacr | ian =e am
4 O Ol O 1

Fic. 2. Female Fellia bicornis (Miiller). a—c, Lateral, ventral and
posterior views of shell. d, First antenna. e, Frontal organ. f, Endop-
odite of second antenna. Scale at bottom right for a-c, at bottom
center for d-f. Scales in mm.

were caught in the upper waters; most were taken over a depth range
of 600-6,000 m, the majority from depths of 1,000-3,000 m.

SUBFAMILY CONCHOECINAE Miiller 1906
Genus Conchoecia Dana 1849

The subfamily Conchoecinae, which originally included all the genera
of Halocyprids except Thaumatocypris, is now, since Poulsen’s (1969)

806 Proceedings of the Biological Society of Washington

revision, restricted to the genus Conchoecia. This genus includes many
species that have been classified into more or less natural groups of
closely related forms.

Spinifera Group

Three species which Miller (1906a) included in this group were
taken at the Barbados station.

Conchoecia spinifera (Claus )

Paraconchoecia spinifera Claus, 1890, p. 14; 1891, p. 65, Pl. X, Figs. 1-7.

Conchoecia spinifera, Miller, 1906a, p. 56, Pl. IX, Figs. 1-10, 14, 15;
1912, p. 69.

Conchoecia spinifera, Deevey, 1968, p. 30, Figs. 8-9.

Specimens of this species were found only in September and October
1963 and in January 1964. C. spinifera was less numerous at the Bar-
bados station than in the Sargasso Sea, where it was found year-round
in the upper 500 m ( Deevey, 1968).

Distribution: 52°N-35°S in the Atlantic, Indian, and Pacific Oceans.

Conchoecia oblonga (Claus)

Paraconchoecia oblonga Claus, 1890, p. 13; 1891, p. 63, Pl. VII, Figs.
10-11, Pl. IX, Figs. 1-14.

Conchoecia oblonga, Miiller, 1906a, p. 58, Pl. IX, Figs. 11-13, 16-25;
1912, p. 69.

Conchoecia oblonga, Skogsberg, 1920, p. 617, Fig. 116.

Conchoecia oblonga, Deevey, 1968, p. 33, Figs. 10, 11.

This was the largest of the dominant species and was found in every
sample, constituting 4.2-10.1 percent of the total numbers of ostracods.

Distribution: 38°N-37°S in the Atlantic and Indian Oceans and Med-
iterranean Sea.

Conchoecia echinata Miller
(Figures 3, 4)

Conchoecia echinata Miiller, 1906a, p. 61, Pl. X, Figs. 14-24; 1906b, p. 3.
Conchoecia notocera Vavra, 1906, p. 58, Pl. 6, Figs. 114-120.
Conchoecia echinata Miller, 1908, p. 67; 1912, p. 70.
Conchoecia echinata, Tes, 1953, p. 268.

Seven females, three males and a number of juvenile specimens of
C. echinata were noted in October 1963 and January, February, April
and May 1964. The females were 1.9-2.0 mm long, the males 1.65-1.7
mm long.

Description: Shell (Fig. 3a, d—g) similar in appearance to that of
C. oblonga, but slightly larger, narrower anteriorly, greatest depth in
the posterior half, with a sharp point at the posterodorsal corner of the
right shell in both sexes. Height of male shell about half the length, of
female shell 42-43 percent of length. The asymmetric glands are in the

North Atlantic ostracods 807

,
alll

Fic. 3. Conchoecia echinata Miiller. a, Lateral view of female shell.
b, Female frontal organ and first antenna. c, Endopodite of female
second antenna (setae and filaments cut off). d, Ventral view of female
shell. e—g, Lateral, posterior, and ventral views of male shell. Scale
at lower left for a, d—g; at lower right for b, c. Scales in mm.

0 O

usual location, the right one clearly defined at the posteroventral corner
of the right shell. The female frontal organ (Fig. 3b) projects well
beyond the tip of the first antenna, the capitulum covered with hairs on
the proximal dorsal half and the ventral surface. The principal seta of the
female first antenna has a row of long hairs anteriorly on the proximal

808 Proceedings of the Biological Society of Washington

0.1

Fic. 4. Male Conchoecia echinata Miller. a, First antenna. b, En-
largement of portion of proximal secondary seta of first antenna. c,
Penis. d, Furca. e, Endopodite of right second antenna (setae and
filaments cut off). £, Left clasping organ. Scale at lower right for e, f;
at center for a, c, d; at upper left for b. Scales in mm.

portion, as in other females of this group, and some spinules distally.
The male principal seta (Fig. 4a) has 17-18 pairs (15, according to
Miiller) of thin teeth directed proximally and two directed distally at

North Atlantic ostracods 809

the distal end of the row of teeth. The distal secondary seta is bare,
but the proximal seta has, three-quarters of the way down its length, a
fan-shaped group of 6-10 spines (Fig. 4b), and more distally a tiny
spine. The male right clasping organ is large and strongly curved, the
left quite small (Fig. 4e, f). The claws on the furca (Fig. 4d) are
almost straight, and this immediately distinguishes this species from
C. oblonga.

Distribution: 31°N-29°S in the Atlantic, Indian, and Pacific Oceans.

Elegans Group
Conchoecia elegans Sars

Conchoecia elegans Sars, 1865, p. 117.

Conchoecia elegans, Skogsberg, 1920, p. 624, Figs. 117, 118.
Conchoecia elegans, Deevey, 1968, p. 40, Fig. 14.

For synonymy, see Skogsberg.

This species was present in all the 400-m samples, and was therefore
somewhat more numerous at this station than in the Sargasso Sea
(Deevey, 1968). As in the Sargasso Sea, all the mature specimens
noted were small, females and males being 1.2-1.3 mm long. Skogsberg
has discussed the extraordinary size range of this species, which is
recorded to have a range of 1.0-2.25 mm in length of mature specimens.

Distribution: 79°58’N-55°S in the Atlantic and Indian Oceans. The
smaller forms of this species have been recorded between 37°N and 24°S
in the Atlantic.

Procera Group
Conchoecia procera Miller
Conchoecia procera Miiller, 1894, p. 228, Pl. 6, Figs. 47, 48, 50-58;

1906a, p. 71, Pl. XIU, Figs. 37-47, Pl. XIV, Figs. 3-6; 1912, p. 72.

Conchoecia procera, Deevey, 1968, p. 45, Figs. 16, 17.

This was one of the dominant species at the Barbados station, and
constituted 7.3-22 percent of the total numbers of ostracods (see Table
1). The females noted were 1.1-1.15 mm long, males 0.95—1.0 mm long.

Distribution: 32°N-37°S in the Atlantic and Indian Oceans and
Mediterranean Sea.

Acuminata Group
Conchoecia acuminata (Claus )
Conchoecetta acuminata Claus, 1890, p. 16; 1891, p. 67, Pls. XIII, XIV.
Conchoecia acuminata, Miiller, 1906a, p. 76, Pl. XV, Figs. 17-23; 1912,
p. 74.
Conchoecia acuminata, Skogsberg, 1931, p. 9.
Conchoecia acuminata, Deevey, 1968, p. 48, Fig. 19.
For further synonymy, see Skogsberg.

This species was present in all 400-m samples. Males were 2.2—2.3 mm,
females 2.9-3.15 mm long.
Distribution: 43°N-37°S in the Atlantic, Indian, and Pacific Oceans.

810 Proceedings of the Biological Society of Washington

Rotundata Group
Conchoecia rotundata Miiller

Conchoecia rotundata Miiller, 1890, p. 275, Pl. XXVIII, Figs. 41-43,
Pl. XXIX, Fig. 44.

Conchoecia rotundata, Deevey, 1968, p. 51, Fig. 20e—j; Fig. 21b, c, e, i-k;
Fig. 22b-e.

C. rotundata was relatively numerous and was taken in every sample,
constituting 1.7-4.2 percent of the total numbers of ostracods. Males
were 0.85-1.1 mm, females 0.85-1.15 mm long.

Distribution: Tropical Pacific Ocean and 15-32°N in the Atlantic
Ocean.

Conchoecia nasotuberculata Miller
(Figures 5, 6)

Conchoecia nasotuberculata Miller, 1906a, p. 83, Pl. XVIII, Figs. 25-
30; 1908, p. 69; 1912, p. 76.
Conchoecia nasotuberculata, Mes, 1953, p. 269.

Only two specimens of this species, a female 0.8 mm long by 0.45 mm
high and a male 0.8 mm long by 0.42 mm high, were taken in the January
1964 sample.

Description: In lateral view the female (Fig. 5a) and the male (Fig.
6a) shells are fairly similar in appearance, narrowed anteriorly, the
anteroventral and posteroventral corners rounded, the height of the
shell slightly greater than half the length. The left asymmetrical gland
protrudes on the left rostrum (Fig. 5f), the right gland is a short distance
below the posterodorsal corner. In ventral and posterior views the
shells of the two sexes are differently shaped. In the male the shoulder
vaults are broadly rounded, the width of the shell is greatest at about
mid-length, and the shell tapers to the posterior end (Fig. 6b, c). The
female shell (Fig. 5b, c) has symmetrically rounded bumps at approx-
imately three-quarters of the shell length, which protrude in ventral and
posterior views, and the rostrum is much narrower than the male’s.

The capitulum of the frontal organ projects beyond the tip of the first
antenna; the female’s (Fig. 5d) is rounded, with hairs or spinules on
the ventral surface, the male’s (Fig. 6d) has relatively long spines
ventrally over the proximal two-thirds of its length and is bent upwards
at the tip. The female principal seta has only a few spinules on the
posterior side of the distal half; the male principal seta has 11-12 pairs
of thin sharp teeth directed proximally. The shapes of the male
clasping organs are shown in Figure 6e-f.

Iles (1953) reported this species abundant at several stations in the
Benguela Current between 22—29°S, and most numerous at depths of
250-500 m.

Distribution: 18°N-—40°S in the Atlantic, Indian Ocean and Mediter-
ranean Sea.

North Atlantic ostracods 811

O O.1 O Ol

Fic. 5. Female Conchoecia nasotuberculata Miiller. a-c, Lateral,
ventral and posterior views of shell. d, Frontal organ and first antenna.
e, Endopodite of second antenna (setae and filaments cut off). f, Inner
view of rostrum. Scale at upper right for a—c; at lower right for e; at
bottom right for f; at bottom left for d. Scales in mm.

Curta Group
Conchoecia curta Lubbock
Conchoecia curta, Miiller 1906a, p. 86, Pl. XXX, Figs. 1-9; 1912, p. 77.
Conchoecia curta, Skogsberg, 1920, p. 661, Fig. 125.
Conchoecia curta, Deevey, 1968, p. 60, Fig. 26.
For synonymy, see Miiller (1906a) and Skogsberg.

This was one of the most abundant species. Except in November,
when it constituted only 1.9 percent of the total numbers of ostracods,

812 Proceedings of the Biological Society of Washington

Bi

(
6

\e

75 6 0.05

Fic. 6. Male Conchoecia nasotuberculata Miiller. a-c, Lateral, ven-
tral, and posterior views of shell. d, Frontal organ and first antenna. e,
Endopodite of right second antenna (setae and filaments cut off). f, Left
clasping organ. Scale at bottom left for a—c; at left center for d; at bot-
tom right for e, f. Scales in mm.

percentages ranged from 12.2-18 percent (see Table 1). Females varied
in length from 0.75—0.85 mm, males 0.75-0.8 mm. Some of the speci-
mens had much more strongly arched shoulder vaults, but no other
differences were noted.

North Atlantic ostracods 813

Distribution: 42°N-37°S in the Atlantic, Indian, and Pacific Oceans
and Mediterranean Sea.
Bispinosa Group
Conchoecia bispinosa Claus

Conchoecia bispinosa Claus, 1890, p. 10; 1891, p. 59, Pl. V, Figs. 1-10,
Pl. VI, Fig. 1, Pl. VIII, Figs. 7, 8.

Conchoecia bispinosa, Skogsberg, 1920, p. 672, Fig. 128.

Conchoecia bispinosa, Deevey, 1968, p. 62, Figs. 27, 28.

This species was recorded from all but the April 1964 sample, and
therefore was somewhat more abundant in these waters than in the
Sargasso Sea, where specimens were taken only occasionally (Deevey,
1968). Females were 1.75-1.95 mm long, males 1.65-1.75 mm long.

Distribution: 42°N-29°S in the Atlantic Ocean.

Conchoecia secernenda Vavra

Conchoecia secernenda Vavra, 1906, p. 59, Pl. VI, Figs. 121-127.
Conchoecia secernenda, Deevey, 1968, p. 65, Figs. 29-31.

C. secernenda was present in every sample but one, and was almost
as numerous at this station as in the Sargasso Sea, where it occurred
year-round in the upper 500 m (Deevey, 1968).

Distribution: 37°N-7°S in the Atlantic Ocean.

Conchoecia atlantica (Lubbock)

Conchoecia atlantica, Miiller, 1906a, p. 92, Pl. V, Figs. 6, 7, Pl. XIX,
Figs. 17-28; 1912, p. 79.

Conchoecia atlantica, Rudyakov, 1962, p. 13, Fig. 9.

Conchoecia atlantica, Deevey, 1968, p. 69, Fig. 32.

For synonymy, see Miiller (1906a).

C. atlantica occurred in every sample, and made up 6 percent of the
total numbers of ostracods in April 1964. This is the largest species
found at this station. Males were 3.4-3.45 mm long, females 3.6—3.65
mm long.

Distribution: 40°N-37°S in the Atlantic, Indian, and Pacific Oceans.

Magna Group
Conchoecia magna Claus

Conchoecia magna Claus, 1874a, p. 6, Pl. I, Fig. 6c, Pl. Il, Figs. 16, 18;
1890, p. 8; 1891, p. 57, Pl. II, Figs. 1-9, Pl. III, Figs. 1, 2.

Conchoecia tetragona Sars, 1887, p. 254, Pl. XI, Figs. 5, 6, Pl. XII,
Figs. 5-9.

Conchoecia magna, Miiller, 1894, p. 228, Pl. V, Figs. 7-12, 16-22, 27-
31, 35-39, 45-52.

Conchoecia magna, Deevey, 1968, p. 77, Figs. 36, 37.

This species was present in all the samples except one.

814 Proceedings of the Biological Society of Washington

Distribution: 52°N-55°S in the Atlantic, Indian, and Pacific Oceans
and Mediterranean Sea.

Conchoecia subarcuata Claus

Conchoecia subarcuata Claus, 1890, p. 9; 1891, p. 58, Pl. III, Figs. 3-9,
Pl. IV.

Conchoecia subarcuata, Miller, 1906a, p. 102, Pl. XXI, Figs. 10-16, 19;
1912, p. 83.

Conchoecia subarcuata, Skogsberg, 1920, p. 695.

Conchoecia subarcuata, Deevey, 1968, p. 86, Figs. 42, 43

For synonymy, see Skogsberg.

A few specimens of this species were taken in September, November,
January and May. Females were 2.0-2.1 mm, males 1.8—1.85 mm long.
Distribution: 37°N-56°S in the Atlantic, Indian, and Pacific Oceans.

Conchoecia spinirostris Claus

Conchoecia spinirostris Claus, 1874, p. 6, Pl. I, Figs. 1, 6a, Pl. Il, Figs.
11, 14, 15; 1890, p. 7; 1891, p. 56, Pl. I, Figs. 1-12.

Conchoecia spinirostris, Miller, 1894, p. 227, Pl. VI, Figs. 1-9, 13.

Conchoecia spinirostris, Skogsberg, 1920, p. 697, Fig. 134.

Conchoecia spinirostris, Deevey, p. 80, Figs. 38, 39.

For further synonymy, see Skogsberg.

This species was present in all the 400-m samples, constituting 2.1-5.1
percent of the total numbers of ostracods. It was also noted in the
surface samples. It was less numerous at the Barbados station than in
the Sargasso Sea, where it was probably the most abundant form in
the upper 500 m ( Deevey, 1968).

Distribution: 45°N-24°S in the Atlantic, the Mediterranean and 33°N
in the Pacific.

Conchoecia porrecta Claus

Conchoecia porrecta Claus, 1890, p. 12; 1891, p. 61, Pl. VII, Figs. 1-13.
Conchoecia porrecta, Deevey, 1968, p. 83, Figs. 40, 41.

C. porrecta was present in every sample, in higher percentages in
October and November, and therefore is a commoner species in the
waters off Barbados than in the Sargasso Sea, where it occurred in-
frequently (Deevey, 1968). Males were 1.3-1.35 mm long, inter-
mediate in size between C. spinirostris and C. parthenoda males, which
they resemble in the shape of the shell. Females were 1.52-1.65 mm
long and are similar in size and appearance to female C. parthenoda, but
differ from the latter species in that the left asymmetrical gland does
not protrude above the dorsal margin of the shell.

Distribution: 41°N-2°N in the Atlantic Ocean.

North Atlantic ostracods 815

Conchoecia parthenoda Miiller
(Figure 7)

Conchoecia parthenoda Miiller, 1906a, p. 78, Pl. XVI, Figs. 24-29.
Conchoecia parthenoda, Deevey, 1968, p. 71, Figs. 33-35.

This species was originally placed in the Obtusata Group, before the
male was described (Deevey, 1968). The male is similar in shape to
all males of the Magna Group, and both males and females appear most
closely related to C. spinirostris and C. porrecta. Conchoecia parthenoda
was a common species at the Barbados station and was found in every
sample, in higher percentages from November to May. Also during the
period from November to May some specimens were present which
differed in that the left asymmetrical gland was located farther forward
on the dorsal margin. Most of these specimens were juveniles, but
males 1.6 mm long were found in the January and May samples. One of
these is illustrated in Figure 7. Aside from being slightly larger (males
of C. parthenoda are 1.35-1.5 mm long), the armature of the principal
seta (Fig. 7g) of these males had several more teeth: 8 pairs of closely
set teeth distally, 6 pairs of alternating teeth, then 12 more widely spaced
teeth proximally, making in profile 31-32 teeth. Also, the left asym-
metrical gland was located farther forward on the dorsal margin, so
that it was 23-24 percent of the total length from the posterodorsal
comer; in the smaller parthenoda males the left asymmetrical gland
is 14-16 percent of the total lengh from the posterodorsal corner.

Distribution: 37°N-30°S in the Atlantic and Indian Oceans.

Conchoecia parvidentata Miller
(Figure 8)

Conchoecia parvidentata Miiller, 1906a, p. 100, Pl. XX, Figs. 11-13;
1908, p. 73; 1912, p. 83.
Conchoecia parvidentata, Skogsberg, 1920, p. 692, Fig. 132.

A single female of this species, 2.55 mm long by 1.23 mm high, was
taken in September 1963.

Description: Shell narrower anteriorly, length slightly more than twice
the height, greatest height in the posterior half, anteroventral and
posteroventral corners rounded (Fig. 8a, b). The asymmetrical glands
are in the usual location, but lateral corner glands are also present, the
one on the right shell just dorsal to the right asymmetrical gland. This
species lacks the gland cells beneath the rostral incisure, which are found
in some members of the Magna Group, and this distinguishes this species
from C. lophura, which is the same size but also has a group of gland
cells on the ventral margin at the posteroventral corner of the left shell.
The frontal organ extends well beyond the first antenna (Fig. 8c); the
capitulum is large and bent downwards from the stem, covered with
spines on the ventral surface and on the dorsal proximal third. The
principal seta of the first antenna has many spinules distally on the

816 Proceedings of the Biological Society of Washington

Fic. 7. Male Conchoecia parthenoda Miller, with left asymmetrical
gland moved farther forward on the dorsal margin. a—c, Lateral, pos-
terior and ventral views of shell. d, Frontal organ and first antenna. e
and f, Endopodites of right and left second antennae (setae and fila-
ments cut off). g, Armature of principal seta of first antenna. Scale
on a for a—c; beside g for g; at left for d; at bottom right for e, f. Scales
in mm.

North Atlantic ostracods 817

Ol

1 a

Fic. 8. Female Conchoecia parvidentata Miiller. a, b, Lateral and
ventral views of female shell. c, Frontal organ and first antenna. d,
Endopodite of second antenna (setae and filaments cut off). Scale at
lower left for a, b; at lower right for c, d. Scales in mm.

posterior surface. According to Skogsberg, the appendages are sim-
ilar to those of C. lophura.

Apparently this species has been recorded only by Miller and Skogs-
berg, and Skogsberg’s specimens were all females. Miiller’s (1906a)
description of the male was brief; he described the armature of the
principal seta as a long double row of small, fine, proximally directed

818 Proceedings of the Biological Society of Washington

9

Fic. 9. Male Conchoecia daphnoides (Claus). a, b, e, Lateral, ven-
tral, and posterior views of shell. c, First antenna. d, Ventral view of
frontal organ. f, Endopodite of left second antenna (setae and filaments
cut off). g, Right clasping organ. Scale at bottom center for a, b, e;
at left for c, d; on f for f, gz. Scales in mm.

teeth, thicker distally than proximally, and hard to distinguish. He
noted that the longer bristle of the basal segment of the endopodite of
the second antenna had strong spinules but lacked the long hairs char-
acteristic of most males of the Magna Group. Miller gave the length of
females as 2.5-2.7 mm, of males as 1.9-2.4 mm.

Distribution: 31°N-48°S in the Atlantic and Indian Oceans.

North Atlantic ostracods 819

Daphnoides Group
Conchoecia daphnoides (Claus )
(Figure 9)

Conchoecilla daphnoides Claus, 1890, p. 17; 1891, p. 68, Pl. XV, Figs.

1-12.
Conchoecia daphnoides, Vavra, 1906, p. 45, Pl. II, Figs. 49-55.
Conchoecia daphnoides var. typica and var. minor, Miiller, 1906a, p.

126, Pl. XXXI, Figs. 1-15.
Conchoecia daphnoides, Skogsberg, 1931, p. 20, Fig. V.
Conchoecia daphnoides, Deevey, 1968, p. 111, Fig. 60.
For further synonymy, see Skogsberg.

Specimens of C. daphnoides were taken in every sample but one.

This species has a wide range in length of mature specimens, fe-
males being 4.2-5.9 mm and males 2.25-3.25 mm long. Although
juvenile stages were present for most of the year in the upper 500 m
of the Sargasso Sea (Deevey, 1968) no mature males were caught. One
male, 2.8 mm long by 0.9 mm high, was taken in the January 1964
sample at the Barbados station.

Description of male: Shell elongate, but less so than female’s, height
approximately one-third length, ventral margin strongly rounded, shoul-
der vaults rounded (Fig. 9a, e), the right asymmetrical gland near the
anteroventral corner beneath the rostrum, the left just below the
posterodorsal corner of the left shell. Sculpture fairly striking, as
illustrated by Miller (1906a, Pl. XXXI, Fig. 1). Three small lateral
gland groups are present, two below the left asymmetrical gland, one
near the posteroventral corner of the left shell. In ventral view (Fig. 9b)
the rostrum is rounded, unlike the female’s which is pointed with the
point of the left rostrum extending well beyond the right. In a ventral
view of the female shell the right posterodorsal corner projects well
beyond the left; in the male the posterodorsal corners are of almost equal
length, the right point slightly longer.

The male frontal organ bends up near the tip, most of the ventral
surface covered with spines (Fig. 9d). The basal segments of the
first antenna are relatively long and slim, and the principal seta is
armed with a long row of approximately 140 pairs of thin fine teeth
directed proximally, but no fine spines directed distally were present.
The secondary setae have only a few spinules near the bend (Fig. 9c).
On the basal segment of the endopodite of the second antenna the
rounded portion that bears the two strong bristles is exceptionally
large, the bristles are sharply bent and covered with spinules. The
clasping organs are shown in Figure 9f, g.

Distribution: Atlantic (60°N-37°S ), Pacific, and Indian Oceans.

The last two species found at the Barbados station have not been
assigned to any group.

820 Proceedings of the Biological Society of Washington

Conchoecia concentrica Miiller
(Figures 10, lla, b)

Conchoecia concentrica Miiller, 1906b, p. 10, Pl. I, Figs. 1-9; 1912,
p. 82.

?Conchoecia pectinata Leveau, 1966, p. 249, Pls. 1, 2.

Conchoecia concentrica, Deevey, 1968, p. 95, Figs. 48-50.

Juvenile specimens of C. concentrica were present in September and
November 1963 and May 1964; one female 1.65 mm long was taken in
April and one male 1.42 mm long in May 1964.

Miiller described this species from three females and one male, and
unfortunately did not figure the female shell. Conchoecia concentrica
varies considerably in the sculpturing of the shell, presumably depending
on the length of time since molting. The specimens found in the
Sargasso Sea (Deevey, 1968) and at the Barbados station appear to differ
from Miiller’s description only in that there is a tiny blunt point at the
posterodorsal corner of the left shell instead of on the right. The
shoulder vaults are swollen and extended laterally, as is evident in anterior
or posterior view, and are blunt-edged in the male (Fig. 1lb), but
sharper-edged in immature specimens and some females (Fig. 10b, d, f),
depending on the extent of the sculpturing. Muller remarked only
“Schulterwulst stark vortretend, stumpfkantig.” As noted previously
(Deevey, 1968), the shoulder vaults of immature specimens and females
“are relatively sharp-edged and may have projecting blunt spines (Fig.
10c-g), evidently prolongations of the sculpturing, which are propor-
tionately larger in smaller individuals or may be lacking or broken off.”
The spines are usually broken off on mature females, so that in anterior
or posterior view the shoulder vaults are blunt-edged (Fig. 10b). It is
possible that the specimens described by Leveau (1966) as Conchoecia
pectinata were immature C. concentrica. Stage IV individuals are 0.8-
0.85 mm long, stage V specimens 1.1-1.3 mm long, the length range of
Leveau’s specimens. The blunt spines on the shoulder vaults of such
immature specimens of C. concentrica (Fig. 10g) are as figured by
Leveau (Pl. 1, Fig. 7) for C. pectinata. Leveau’s figures indicate that he
was describing immature specimens. His drawing of the furca of C.
pectinata, for example, shows only 7 claws and an unpaired bristle on
the furca. The number and lengths of the claws are similar in 1.1-1.3
mm long juveniles of C. concentrica.

In some respects C. concentrica resembles members of the Bispinosa
Group. The capitulum of the male frontal organ is bent upwards and
spined as in C. bispinosa; also the proximal secondary seta of the male
first antenna has a pad or callous, such as is found in the C. bispinosa
male. However, one of the two setae of the second segment of the
endopodite of the male second antenna is not strikingly long, and the
female lacks the extra bristle on this segment. Conchoecia concentrica
differs also in having lateral corner glands near the posteroventral corners

North Atlantic ostracods 821

Fic. 10. Female Conchoecia concentrica Miller. a, b, Lateral and
posterior views of a specimen from the Barbados station. c—f, Lateral,
posterior, ventral and anterior views of specimen from the Sargasso Sea
still retaining spines on shoulder vaults. g, Enlargement of spine. Scale
at lower left for a—-f; at lower center for g. Scales in mm.

of both shells. Skogsberg (1920) believed that C. concentrica might
be related to C. serrulata,

Distribution: As previously known, the distribution was the Malay
Archipelago and 38°N-32°N in the Atlantic Ocean; the Barbados speci-
mens extend the range to 15°N.

822 Proceedings of the Biological Society of Washington

Fic. 11. a, b, Lateral and posterior views of male Conchoecia con-
centrica Miiller. c—e, Lateral, ventral, and posterior views of juvenile
Conchoecia sp. Scale for a—e, in mm.

Conchoecia sp.
(Figure 1lc-e)

Conchoecia sp., Deevey, 1968, p. 114, Fig. 61.

Five juvenile specimens of a species not yet named due to lack of
mature specimens were found in September 1963 and May 1964. These
juveniles were 0.6, 1.15, 1.2, 1.6, and 1.65 mm long. This species
resembles C. concentrica in the shape of the shell, but is larger at maturity
and of slim build, whereas C. concentrica is a plump species, the body
always filling the shell. The shell appears to lack sculpturing, but faint
lines may be seen, running anteroposteriorly in a pattern similar to the

North Atlantic ostracods 823

sculpturing of the C. concentrica shell. The asymmetrical glands and
lateral corner glands are situated as in C. concentrica. The postero-
dorsal corners of both shells are rather bluntly rounded and of equal
size, neither the right nor the left shell produced into a definite point.
Lateral, posterior, and ventral views of the 1.2 mm juvenile shell are
shown in Figure 1lc—e.

Distribution: 32°N-15°N in the Atlantic Ocean.

LITERATURE CITED

Beers, J. R., D. M. STEVEN, AND J. B. Lewis. 1965. Primary produc-
tivity in the tropical North Atlantic off Barbados and the
Caribbean Sea off Jamaica. Bermuda Biological Station
Final Report to ONR, March, 1965.
. 1968. Primary productivity in the Caribbean Sea off Jamaica
and the tropical North Atlantic off Barbados. Bull. Mar.
Sci., 18: 86-104.
Ciaus, C. 1874. Die Familie der Halocypriden. Schriften Zoolog.
Inhalts. H. I, Wien.
. 1890. Die Gattungen und Arten der mediterranen und
atlantischen Halocypriden. Zool. Inst. Univ. Wien, Arb. IX:
1-34.
1891. Die MHalocypriden des Atlantischen Oceans und
Mittelmeeres. Wien, Alfred Holder. 81 p., 26 pls.
DEEvEY, G. B. 1968. Pelagic ostracods of the Sargasso Sea off Ber-
muda. Peabody Mus. Nat. Hist., Yale, Bull. 26, 125 p., 65
figs.
ILes, E. J. 1953. A preliminary report on the Ostracoda of the Ben-
guela Current. Discovery Rept. XXVI: 259-280.
Leveau, M. 1966. Conchoecia pectinata: nouvelle espece d’ostracode
pelagique. Rec. Trav. Sta. mar. Endoume. Bull. 40, Fasc. 56:
249-952. Pls. 1-2.
Muuier, G. W. 1890. Ueber Halocypriden. Zool. Jahrb. Abt. Syst.,
Geog. Biol. V (2): 253-280, Pls. 28-29.
. 1894. Die Ostracoden des Golfes von Neapel. Fauna und
Flora des Golfes von Neapel. Monogr. 21: 1404. Pls. 1-40.
. 1906a. Ostracoda. Wissensch. Ergeb. d. Deutschen Tief-
see-Expedition auf dem Dampfer “Valdivia” 1898-1899. 8:
1-154, Pls. V-XXXV.
1906b. Die Ostracoden der Siboga-Expedition, Uitkomsten
of Zoologisch, Botanisch, Oceanographisch en Geologisch
Gebied versameld in Nederlandsch Oost-India 1899-1900.
Siboga-Expeditie XXX: 1-40, Pls. I-IX.
1908. Die Ostracoden der Deutschen Siidpolar-Expedition
1901-1903. Deutsche Siidpolar-Expedition 1901-1903, X,
Zool. II; 53-181, Pls. IV—XIX.
— ——. 1912. Ostracoda. In Das Tierreich, Lief. 31, 434 p. Berlin.
Pouusen, E. M. 1969. Ostracoda-Myodocopa Part IIIA Halocypri-

824 Proceedings of the Biological Society of Washington

formes-Thaumatocypridae and Halocypridae. Dana-Report
No. 75. 100 p., 40 figs.

Rupyaxov, Yu. A. 1962. Ostracoda Myodocopa (family Halocyp-
ridae) from the N. W. Pacific Ocean. Trudy Inst. Okeanol.,
58: 172-201. 14 figs.

Sars, G. O. 1865. Oversight of Norges marine Ostracoder. Sectio 2.
Myodocopa. Forh. Videnskabs-Selskabet, Christiania 1865:
99-120.
1887. Nye Bidrag til Kundskaben om Middelhavets Inverte-
bratfauna. IV. Ostracoda Mediterranea. Arch, Math. Naturv.
XII: 173-324, Pls. I-XX.

SxocsBERG, T. 1920. Studies on marine ostracods Part I (Cypridinids,
Halocyprids and Polycopids). Zool. Bidr. Uppsala. Suppl. I,
784 p., 153 figs.
1931. Ostracods. Rep. Sci. Res. “Michael Sars” North At-
lantic Deep-Sea Expedition 1910. V: 3-26, 5 figs.

Vavra, V. 1906. Die Ostracoden (Halocypriden und Cypridiniden)
der Plankton-Expedition. Ergebn. Plankton-Exp., II, G, g,
76 p., 8 pls.

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63, pp. 825-828 5 February 1970

PROCEEDINGS
OF THE

BIOLOGICAL SOCIETY OF WASHINGTON

A NEW SPECIES OF EUPERA (MOLLUSCA; _
PELECYPODA) FROM HAITI q

By Micuet A. KLAPPENBACH
Museo Nacional de Historia Natural
Montevideo, Uruguay

The genus Eupera Bourguignat, 1854, is represented in the
Greater Antilles by the following species: E. cubensis (Prime,
1865) from Cuba; E. portoricensis (Prime, 1863) and E.
parvula (Prime, 1865) from Puerto Rico and E. veatleyi (C. B.
Adams, 1849) from Jamaica, which to date has not been cited
from Hispaniola.

While recently checking material of this genus in the col-
lections of the U.S. National Museum, I came across an
undetermined lot consisting of a large number of isolated
valves (possibly inarticulated when collected) found in the
southwest of the island, near Los Cayes, Dept. du Sud, Haiti.
Having studied this material, I arrived at the conclusion that
it represented a new species, which I describe as follows:

Eupera haitiensis new species
Figures 1-3

Description: (Holotype, left valve). (Fig. 1). Shell of medium size
for the genus; subovate, short and very deep for its size; margins smooth,
slightly arched, united in a regular curve that does not present any
particular characteristic. The anterior margin much shorter than the
posterior; the inferior, the longest.

Umbo large, with a forward and inward projection and situated on a
level with the first anterior third of the shell, while projecting conspic-
uously above the superior margin (Fig. 2). Cardinal tooth, small and
simple. In its lateral aspect it has the appearance of a very small, short
and straight lamella, obliquely located on the curved hinge, beneath
and to the rear of the umbo. In its inferior aspect (Fig. 3), it has the
appearance of a small, truncate cone, the anterior face of which con-
tinues slightly downwards, while the posterior, short side, terminates
abruptly and almost vertically.

63—Proc. Brot. Soc. WaAsH., Vou. 82, 1970 (825)

eel

826 Proceedings of the Biological Society of Washington

\

“seer

A new pelecypod 827

This tooth is separated from the anterior margin by a small and not
very deep parallel groove that engages the cardinal of the opposite valve.

On its anterior side an articular surface is present where the interior
face of the right cardinal rests. Lateral teeth, simple; the anterior
tooth with a short but robust base, appears from below as a higher cone
than that of the cardinal. It is separated from the exterior margin by
a small groove and presents on its superior and inferior faces a sharply
defined articular surface.

The posterior lateral, lower and more extended, presents in its inferior
aspect a conical profile, low but very regular. It, too, is separated from
the external margin by an extended groove.

As in the other lateral, it presents on its superior and inferior faces a
clearly defined articular surface.

The ligament, fine and elongate, extends from the umbo as far as the
origin of the posterior lateral. It is limited inferiorly by a thin but quite
perceptible ridge, parallel to the superior margin.

The internal surface, brilliant beyond the pallial line and in the im-
pressions of the adductors, but opaque in the rest of the shell.

Color white, but splotchy, with characteristic purplish-brown spots,
clotted and clustered, although limited in expansion by the pallial line.

The external surface, wanting in periostracum, presents a dullish
white color with very fine concentric lines of growth alternating with
larger, thick, rough and irregular ridges, of which it is possible to discern
four.

Holotype: U.S.N.M. 404968. Left valve. Collected by C. R. Orcutt.
Date: 17 May 1929

Type-Locality: O’Shell Sugar Plantation, near Los Cayes, Dept. du
Sud, Haiti

Measurements: (in mm) 5.0 x 4.0

Paratypes: U.S.N.M. 679533. Same data as the holotype
M.N.H.N. Montevideo. 1328. Same data as the holotype
U.S.N.M. 440232. Los Cayes, Dept. du Sud, Haiti (Or-
cutt! )
U.S.N.M. 439871. Bizoton, Dept. de POuest, Haiti (Or-
cutt! )

Remarks on Paratypes (Right Valve): Cardinal weaker. It appears as a
small oblique fold that in its inferior aspect does not extend beyond the
superior margin of the shell. On its inferior face, however, it presents
a very small articular surface that is separated from the exterior margin
by a shallow cavity.

<

Fics. 1-3. Eupera haitiensis n. sp. (Holotype, U.S.N.M. 404968).
Fig. 1—Internal view. Fig. 2—External view. Fig. 3—The hinge.

828 Proceedings of the Biological Society of Washington

The anterior laterals, though solid, are low and very short (the
inferior the larger), and are separated from one another by a relatively
deep cavity.

The posterior laterals (the inferior the larger) are also low but more
extended than the anterior, and are separated by a narrow groove that is
not so deep but longer than that of the anterior.

The inner faces of the laterals (both anterior and posterior) are finely
granulate (as shown under magnification). The zones where the peri-
ostracum is still present are of a greenish-gray color and are not very
brilliant. Over and above this, they exhibit the concentric lamellae
already described in other species of the genus.

In some of the young specimens the periostracum presents a light-
brownish color. Practically all the specimens show the thick and rough
concentric ridges that were observed in the holotype; the number vari-
able but as many as 15.

The depth of the valves and the strong teeth as compared with these
features in other species of the genus is striking. The shape of the shell
is remarkably constant in the whole lot.

Discussion: Compared with E. cubensis, the new species that we
describe here may be easily distinguished by its different shape, charac-
terized by its shorter posterior margin and more developed umbo.

In E. cubensis, the superior margin is higher posteriorly than the umbo,
it being the highest part of the shell.

In E. haitiensis, on the other hand, the umbo is the highest part of the
shell and definitely exceeds the superior margin. The cardinal tooth of
the left valve is lower and more robust in E. haitiensis, while in E.
cubensis it is higher and weaker. Nevertheless, the laterals are more
elongated in this last species. The same occurs in the case of the other
forms, E. veatleyi, E. portoricensis and E. parvula, of the area.

I may add that the position of the umbo is more central in E. haitiensis
than in the other above-mentioned species. The same applies to E.
bahamensis Clench, with which I have also compared E. haitiensis.

LITERATURE CITED

Apams, C. B. 1849. Descriptions of supposed new species of fresh
water shells which inhabit Jamaica, Contributions to Con-
chology, 1:42-45.

CiencH, W. J. 1938. Origin of the land and freshwater mollusk fauna
of the Bahamas, with a list of the species occurring on Cat
and Little San Salvador Islands, Bull. Mus. Comp. Zool.
Harvard, 80 (14): 481-541, pls. 1-3, text figs. 1-2.

Prme, T. 1861. Descriptions of three new species of Mollusca of the
Genus Sphaerium, Proc. Acad. Nat. Sci. Philadelphia, 13:
414-415.

1865. Monograph of the American Corbiculadae. (Recent
and Fossils.) Smithsonian Miscellaneous Collections, 7 (5):
1-80, text figs. 86.

73

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o. 64, pp. 829-836 5 February 1970

PROCEEDINGS
OF THE

BIOLOGICAL SOCIETY OF WASHINGTON

A NEW CRAYFISH OF THE GENUS FALLICAMBARUS
FROM TENNESSEE (DECAPODA, ASTACIDAE)

By Horton H. Hosss, Jr. AND JosePH F, FirzPATRICK, JR.

Smithsonian Institution and
Randolph-Macon Woman's College

Among the many undescribed crayfishes known to occur in
Tennessee, this new member of the burrowing genus Fallicam-
barus was dug from comparatively simple shallow burrows
constructed in sandy soil near a small tributary to the Hatchie
River in McNairy County. Its range lies along the edge of the
boundary of that outlined for the genus by Hobbs (1969: 124),
extending “from southern Ontario, Michigan and Illinois south-
ward to Texas and across western Tennessee to southwestern
Georgia; east of the Appalachians it extends from Maryland
to South Carolina.”

We are grateful to H. H. Hobbs III, Daniel J. Peters, and Dr.
Jean E. Pugh for their assistance in obtaining the 24 specimens
on which this description is based.

Fallicambarus hortoni new species

Diagnosis: Body pigmented, eyes well-developed. Rostrum depressed,
acuminate, and devoid of marginal spines or tubercles. Areola obliter-
ated or linear, its projected extent comprising 36.0 to 38.2 per cent
of entire length of carapace. Cervical spines or tubercles lacking. Sub-
orbital angle weak, obtuse. Postorbital ridges terminating cephalically
with or without very small tubercles. Antennal scale 2.3 to 2.5 times
longer than broad, broadest distal to midlength. Chela with two rows of
tubercles on mesial surface of palm; lateral margin of chela costate and
both fingers with well-defined longitudinal ridge on upper surface; dactyl
with distinct emargination. First sinistral pleopod (Figs. 1, 5, 8) of first
form male with corneous central projection recurved at approximately 90
degrees and strongly deflected dextrally, scarcely tapering distally,
broadly truncate with subapical notch lacking, or perhaps represented by
shallow emargination; non-corneous mesial process only slightly tapering

; .64—Proc. Brot. Soc. Wasu., Vou. 82, 1970 (829)
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830 Proceedings of the Biological Society of Washington

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rite ae

Fics. 1-12. Fallicambarus hortoni new species (pubescence removed
from all structures illustrated except for Figs. 1, 8, 9, 10). 1, Mesial
view of first pleopod of holotype. 2, Mesial view of first pleopod of
morphotype. 38, Dorsal view of carapace of holotype. 4, Lateral view
or first pleopod of morphotype. 5, lateral view of first pleopod of

A new Fallicambarus 831

distally, eminence on morphological cephalic border almost at tip of proc-
ess and overlapping central projection laterally with small constricted dis-
tal portion extending slightly beyond tip of central projection. Annulus
ventralis (Fig. 6) immovable, approximately 1.6 times broader than
long, deeply excavate dextrally, and with conspicuous tongue sloping
cephalosinistrally from elevated caudal wall. Color olive brown with
irregular dark and light markings and sometimes with median longitudinal
pale olive-tan stripe.

Holotypic Male, Form I: Body subovate, slightly compressed. Abdo-
men narrower than cephalothorax (11.0 and 12.6 mm). Carapace
broader than depth at caudodorsal margin of cervical groove (12.6 and
10.8 mm). Areola linear and constituting 37.5 per cent of entire length
of carapace; cephalic section of carapace 1.7 times longer than areola.
Rostrum depressed, acuminate, excavate dorsally with converging thick-
ened margins; marginal spines or tubercles lacking; upper surface with
conspicuous, deep punctations, submarginal rows of setiferous puncta-
tions becoming progressively shallower toward apex; acumen slightly
upturned and reaching slightly beyond base of ultimate podomere of
peduncle of antennule; subrostral ridges visible in dorsal aspect almost
to midlength of rostrum. Postorbital ridges strong, grooved dorso-
laterally, and terminating cephalically without tubercles. Suborbital angle
obtuse and weak. Branchiostegal spine reduced to angle. Carapace
with many prominent setiferous punctations dorsally and dorsolaterally,
but less dense cephalolaterally, and with setiferous squamous tubercles on
lateral branchiostegal region, anteriormost forming row along caudo-
ventral margin of cervical groove. Cervical spines or tubercles lacking.
Abdomen longer than carapace (26.2 and 24.5 mm). Cephalic portion
of telson with two strong spines in each caudolateral corner, mesial ones
movable. Uropods with distolateral margin and distal end of submedian
ridge of outer ramus with strong acute spines, that on ridge almost
reaching distal margin of ramus; distal portion of proximal segment of
inner ramus with row of strong acute spines; basal segment of uropods
with strong mesiodistal acute spine overlapping base of outer ramus;
smaller acute laterodistal spine also present.

Epistome (Fig. 7) about 1.5 times longer than broad, subtriangular,
nearly plane, and bearing small apical notch and shallow caudomedian
fovea. Antennules of usual form with well-developed small spine on
mesioventral margin slightly distal to midlength. Antennae broken
but probably reaching beyond midlength of abdomen. Antennal scale
(Fig. 12) 2.3 times longer than broad, broadest distal to midlength with

<

holotype. 6, Annulus ventralis of allotype. 7, Epistome of holotype.
8, Caudal view of first pleopods of paratypic male, form I. 9, Bases of
third, fourth, and fifth pereiopods of holotype. 10, Dorsal view of distal
podomeres of cheliped of holotype. 11, Lateral view of carapace of
holotype. 12, Antennal scale of holotype.

832 Proceedings of the Biological Society of Washington

widest lamellar area approximately 1.6 times width of thickened lateral
portion, latter terminating in strong acute spine.

Right chela (Fig. 10) strongly depressed with palm inflated and bear-
ing scattered punctations proximally, becoming more numerous distally,
and particularly crowded and conspicuous at base of immovable finger.
Inner margin of palm with mesial row of eight tubercles subtended
dorsolaterally by row of eight with single tubercle immediately lateral
to distal one in second-mentioned row. Ventral surface of palm with
widely scattered setiferous punctations and with prominent tubercle at
base of dactyl. Fingers only slightly gaping. Opposable margin of
immovable finger with two prominent tubercles in proximal half and
single row of minute denticles extending distally from distal tubercle;
single large tubercle in proximal portion of distal half below row of
denticles; upper surface with strong submedian ridge flanked by deep
setiferous punctations, and less conspicuous ridge mesially; lateral margin
of finger strongly costate, ridge extending proximally onto distal portion
of palm; ventral surface with distinct row of setiferous punctations along
lateral margin and heavily bearded in basal eighth of mesioventral por-
tion. Opposable margin of dactyl distinctly excised, with two promi-
nent tubercles in basal half of excision and one large tubercle at its
distal end, latter tubercle followed distally by single row of minute
denticles interrupted by two small tubercles; dorsal surface with strong
submedian longitudinal ridge flanked by setiferous punctations, and six
small tubercles flanking mesial row of punctations; mesial margin with
row of 13 tubercles decreasing in size distally and extending almost entire
length of dactyl.

Carpus of cheliped longer than broad with deep submedian longitudinal
furrow; dorsal surface with scattered setiferous punctations; mesial sur-
face with prominent acute spine slightly distal to midlength and with
row of four much smaller spiniform tubercles proximal to it, latter flanked
above and below by irregular cluster of small tubercles; ventral latero-
distal and mesiodistal extremities with acute spines.

Merus of cheliped with upper surface sparsely punctate and bearing
two obscure tubercles distally; mesial and lateral surfaces also sparsely
punctate; ventral surface with mesial row of 15 heavy acute tubercles
and lateral one of two acute tubercles in middle third... Ischium with
single very small tubercle on mesial margin proximal to midlength.

Hooks on ischia of third pereiopods only (Fig. 9); hooks simple, ex-
tending proximally beyond distal margin of basis and not opposed by
tubercle on latter. Coxa of fourth pereiopod with prominent longitu-
dinally oriented furrow with conspicuous rounded elevation mesial to it
and obliquely directed boss caudomesially, boss excavate anteroventrally.
Coxa of fifth pereiopod with only slight caudomesial elevation.

Sternum between bases of third, fourth, and fifth pereiopods deep
and with conspicuous setal mat extending mesioventrally and covering
first pleopods.

First pleopods (Figs. 1, 5, 8) symmetrical basally but markedly asym-

A new Fallicambarus 833

metrically disposed distally with central projection of sinistral member
of pair directed caudodextrally across median line of body, displacing
corresponding element of dextral pleopod laterally (Fig. 8). Pleopods
reaching base of coxae of third pereiopods when abdomen is flexed and
terminating in two parts bent caudally at approximately right angles to
main axis of shaft of appendage (see diagnosis for description).

Morphotypic Male, Form II: Differs from holotype in following re-
spects: rostrum with apical tubercle much reduced, not upturned, and
not reaching base of ultimate podomere of peduncle of antennule;
epistome subtriangular with rounded apex; right chela apparently regen-
erated but mesial margin of palm of left chela with most mesial row of
only 7 tubercles; distal tubercle on opposable margin of immovable
finger much reduced; mesial margin of dactyl with row of 12 tubercles
and with row of 5 tubercles immediately lateral to it; carpus of chela
not so distinctly longer than broad, and major spine on mesial surface
surrounded by irregularly arranged tubercles; upper distal surface of
merus with three small tubercles, ventrolateral margin with three spines;
hooks on ischia of third pereiopods and ornamentation of coxa of
fourth not so strongly developed.

First pleopod (Figs. 2, 4) with no corneus elements; central pro-
jection broadly rounded with faint indication of emarginations apically;
mesial process also broadly rounded and directed somewhat laterally
with eminence on morphological cephalic border much reduced; basal
“segment” of pleopod delimited by suture.

Allotypic Female: Differs from holotype in following respects: abdo-
men and cephalothorax subequal in width; mesial row of six tubercles on
inner margin of palm of chela subtended dorsolaterally by row of
seven; opposable margin of dactyl with row of four tubercles distal
to large tubercle at distal end of excision; mesial surface of dactyl
with lateral row of only five tubercles; carpus with two spinous tubercles
aligned between strong mesial spine and mesiodistal ventral spine; dorsal
surface of merus with irregular row of five tubercles distally, ventro-
mesial margin with row of 14 and ventrolateral margin with row of three.

Sternum between last three pairs of pereiopods deep. Annulus ven-
tralis (Fig. 6) immovable but with distinct groove between it and
sternum immediately cephalic to it, about 1.6 times broader than long,
and with caudal margin markedly elevated (ventrally); deep sinus
originating near median line caudal to midlength, extending approxi-
mately one-fourth width of annulus, and recurving rather suddenly
caudodextrad to cut caudal margin just sinistral to median line; tongue
sloping cephalosinistrally from caudal wall to dip below transverse portion
of sinus; usual median trough displaced dextrally, extending caudo-
dextrally from midcephalic margin of annulus to base of elevated caudo-
dextral wall, becoming broader and deeper caudally. Sternite immediately
caudal to annulus broadly triangular, about 2.4 times broader than long,
not highly elevated but with highest portion centrally situated.

Type-locality: Low area along a roadside ditch leading into a small

834 Proceedings of the Biological Society of Washington

Measurements: As follows (in mm):

Holotype Allotype Morphotype

Carapace

Height 10.8 LOW 7.6

Length 24.5 25.9 19.2

Width 12.6 12.5 9.3
Rostrum

Length 6.4 6.0 4.5

Width 4.1 4.3 3.4
Areola length 9.2 9.6 ol
Chela, right

Length of outer margin 19.0 15.9 10.9:

Length of inner margin of palm 4.7 4.5 2.9

Width of palm 7.9 7.0 Dat

Length of dactyl 13.1 11.0 7.8

tributary of Cypress Creek, 7.5 miles east of the Hardeman County line
on State Route 57 (Hatchie River drainage), McNairy County, Tennes-
see. The animals were dug from burrows (see introductory paragraph),
some of which were provided with chimneys similar to those constructed
by Cambarus d. diogenes Girard, 1852. Vegetation in the area consisted
of several species of grasses and Compositae and a member of the genus
Viola. Salix nigra was along the ditch, and slightly more distant were
trees belonging to the genera Acer, Liquidambar, and Liriodendron.

Disposition of Types: The holotypic male, form I, the morphotypic
male, form II, and the allotypic female are deposited in the United States
National Museum (nos. 129895, 129896, and 129897, respectively ). Para-
types also located at the USNM include 361, 3¢II, 129, and in the
collection of the junior author are 1¢1, 19, and a damaged ¢II. All
were collected from the type-locality on 20 May 1969 by those mentioned
above and the authors.

Color Notes: Carapace and abdomen drab olive-brown with olive
black and pale olive tan mottlings; branchiostegal areas fading ventrally.
Some specimens with discreet pattern consisting of dorsomedian longi-
tudinal light stripe extending from base of telson cephalically to cervical
groove, flanked laterally by very dark irregularly margined dark stripes
from telson to at least midlength of areola. Caudodorsal margin of pleura
of abdomen with pale spot. Upper surface of distal podomeres of chelae
very dark, somewhat paler below with tips of fingers reddish orange.
Proximal podomeres of remaining pereiopods pale, merus and more
distal podomeres dark, darker dorsally than ventrally. Lateral portions
of cephalic section and most of caudal section of telson dark as are
inner ramus and distal segment of outer ramus of uropods and mesiodistal
end of uropodal peduncle.

A new Fallicambarus 835

Range and Crayfish Associates: Fallicambarus hortoni is known only
from the type-locality. No other primary burrowing species were en-
countered in the immediate vicinity, but Cambarus striatus Hay, 1902,
was collected from the adjacent creek as were Procambarus ablusus Penn,
1963, and an undescribed species of the genus Orconectes. Nearby
localities yielded specimens of C. d. diogenes Girard, 1852, and what
appears to be C. d. ludovicianus Faxon, 1914. Further collections in the
area are needed to determine the precise degree of overlap (ecological
and geographical) which exists between these primary burrowing cray-
fishes.

Variations: Little variation was observed in the type series beyond that
usually encountered in a crayfish population. In one first form male,
however, the branchiocardiac grooves are sharply arced so that the
areola, although obliterated, is not linear, and one punctation is evident
anterior and posterior to the obliterated portion.

Relationships: Fallicambarus hortoni is the first species to be assigned
originally to the genus Fallicambarus, Hobbs, 1969. Of the eight species
previously recognized, F. hortoni is probably more closely allied to F.
byersi (Hobbs, 1941) and F. oryktes (Penn and Marlow, 1959) than to
any of the others. The latter two occur in the lower coastal plain between
the Choctawhatchee River, Florida and the Pontchartrain Basin in Louisi-
ana. The kinship is most clearly seen in the similarities of the first
pleopods and annulus ventralis. More distantly, it is related to F. fodiens
(Cottle, 1863), F. hedgpethi (Hobbs, 1948), and F. uhleri (Faxon,
1884) which species occupy the northwestern, southwestern, and eastern
limits of the range of the genus. The peculiar structure of the distal
portion of the first pleopod of the first form male, however, readily
distinguishes it from any other crayfish.

Etymology: We are pleased to name this crayfish in honor of Horton
H. Hobbs II, who, in his work on the entocytherid ostracods associated
with primary burrowing crayfishes, has added greatly to our knowledge of
the ranges of many of the burrowing species.

LITERATURE CITED

Corte, T. J. 1863. On the two species of Astacus found in upper
Canada. Canad. Journ. Industry, Sci., and Arts (45, n.s.):
216-219.

Faxon, WALTER. 1884. Descriptions of new species of Cambarus, to
which is added a synonymical list of the known species of
Cambarus and Astacus. Proc. Amer. Acad. Arts and Sci., 20:
107-158.
1914. Notes on the crayfishes in the United States National
Museum and the Museum of Comparative Zoology with de-
scriptions of new species and subspecies to which is appended
a catalogue of the known species and subspecies. Mem. Mus.
Comp. Zool., Harvard Coll., 40 (8): 351-427, 13 pls.

Gmarp, CHARLES. 1852. A revision of the North American Astaci,

836 Proceedings of the Biological Society of Washington

with observations on their habits and geographical distri-
bution. Proc. Acad. Nat. Sci., Philadelphia, 6: 87-91.

Hay, WILLIAM Perry. 1902. Observations on the crustacean fauna of
Nickajack Cave, Tennessee, and vicinity. Proc. U.S. Nat.
Mus., 25 (1292): 417-439, 8 figs.

Hosss, Horton H., Jr. 1941. Three new Florida crayfishes of the
subgenus Cambarus (Decapoda, Astacidae). Amer. Midl.
Nat., 26 (1): 110-121, 2 pls.
1948. A new crayfish of the genus Cambarus from Texas,
with notes on the distribution of Cambarus fodiens (Cottle).
Proc. U. S. Nat. Mus., 98 (3230): 223-231, 1 fig.
1969. On the distribution and phylogeny of the crayfish
genus Cambarus. In Holt, Perry C., Richard L. Hoffman, and
C. Willard Hart, Jr., editors. The distributional history of
the biota of the Southern Appalachians, Part I: Inverte-
brates. Va. Polytechnic Institute, Research Division Mono-
graph 1: 93-178, 20 figs.

PENN, GeorcE Henry, Jr. 1963. A new crawfish from the Hatchie
River in Mississippi and Tennessee (Decopoda, Astacidae).
Proc. Biol. Soc. Wash., 76: 121-126, 10 figs.

AND Guy MarLow. 1959. The genus Cambarus in Louisiana.

Amer. Midl. Nat., 61 (1): 191-203, 14 figs.

Vol. 82, No. 65, pp. 837-842 5 February 1970

PROCEEDINGS
OF THE

BIOLOGICAL SOCIETY OF WASHINGTON

FAMILIAL TAXA WITHIN THE CAPRELLIDEA
(CRUSTACEA: AMPHIPODA )

By Joun C. McCain
Smithsonian Institution, Washington, D. C.

Historically the Caprellidea have been divided into two
families, the Cyamidae (whale lice) and the Caprellidae
(skeleton shrimps). Due primarily to the recent discovery of
Caprogammarus, an intermediate form between the caprel-
lidean and gammaridean stocks, questions have been raised
about this classical division.

Vassilenko (1968) proposed a new family, Paracercopidae
(invalid name), for the genus Cercops and the following four
subfamilies for the remaining genera of Caprellidae:

Phtisicinae Vassilenko—Phtisica, Paraproto, Protogeton, Pro-
toplesius, Pseudoproto, Protomima, Metaproto.

Dodecadinae Vassilenko—Dodecas, Dodecasella, Caprellina,
Hircella, Pseudocaprellina, Liriarchus, Aeginoides.

Aeginellinae Vassilenko—Aeginella, Aeginina, Thorina, Pro-
tellina, Proaeginina, Parvipalpus, Parvipalpina.

Caprellinae Dana—Aciconula, Caprella, Caprellinoides,
Deutella, Eugastraulax, Eupariambus, Hemiaegina, Liropus,
Luconacia, Mayerella, Metaprotella, Monoliropus, Noculacia,
Orthoprotella, Paedaridium, Paracaprella, Paradeutella, Pari-
ambus, Paraprotella, Pedoculina, Piperella, Proliropus, Propo-
dalirius, Protella, Protellopsis, Pseudaeginella, Pseudolirius,
Pseudoprotella, Triantella, Triliropus, Triperopus, Tritella.

McCain (1968, pp. 107-112) discussed the relationship be-
tween the suborders Gammaridea and Caprellidea and stated
that a revision of familial taxa is necessary in the Caprellidea
but that the mouthparts of many Caprellidae were too poorly
known to allow such a revision at that time. Since then, numer-
ous species have been examined. The mandible shows several

65—Proc. Biot. Soc. Wasu., Vou. 82, 1970 (837)

838 Proceedings of the Biological Society of Washington

trends which roughly coincide with Vassilenko’s conception of
the subfamilies based primarily on degrees of pereopod reduc-
tion.

Four distinct groups of mandible types are evident in the
Caprellidea as follows:

Group I. Mandible lacking molar, bearing numerous spines
and accessory plates, and bearing a mandibular palp.

Group II. Mandible with molar, bearing 2 or 3 spines and a
lacinia mobilis and incisor, and with a mandibular palp.

Group III. Mandible with molar, bearing 2 or 3 spines and
a typical lacinia mobilis and incisor, and lacking a mandibular
palp.

Group IV. Mandible without molar or palp.

Group I includes Vassilenko’s concept of the Phtisicinae and
Dodecadinae and Group II the Aeginellinae. The Caprellinae,
however, appear to be a heterogeneous assemblage of mandi-
ble types.

McCain (1968, p. 3) stated that the mouthparts undoubtedly
reflect feeding habits of caprellids and thereby, to some extent,
their niche. Since we lack a fossil record of the Caprellidea, the
choice of mandible types as conservative characters seems
justified. Lacking other obvious conservative characters, the
use of mandible types to characterize higher taxa should be
much more valid than the use of reduced segmentation of
vestigial appendages.

I, therefore, propose the following familial classification of
the Caprellidea:

PHTISICIDAE VASSILENKO, 1968, emend.

Mandible lacking molar, with mandibular palp (Group I);
pereopods 3-5 fully segmented or reduced; gills on pereonites
2-4, rarely 3-4; abdomen of single reduced article. Two sub-
families:

Phtisicinae Vassilenko, 1968, emend.

Pereopods 3-5 fully segmented, pereopod 5 rarely reduced.
Nine genera: Hemiproto, Metaproto, Paraproto, Phtisica, Pro-
togeton, Protomima, Protoplesius, Pseudoproto, Pseudopro-
tomima.

Caprellid families 839

Dodecadinae Vassilenko, 1968, emend.

Pereopeds 3-5 variably reduced, pereopod 5 of less than 4
articles except for 1 genus. Twelve genera: Aeginoides, Cap-
rellina, Caprellinoides, Dodecas, Dodecasella, ? Fallotritella,
Hircella, Liriarchus, Peadaridium, Pereotripus, Prellicana,
Pseudocaprellina.

AEGINELLIDAE VASSILENKO, 1968, emend.

Mandible with molar and mandibular palp (Group II);
pereopods 3-4 considerably reduced or absent, pereopod 5
occasionally reduced; gills on pereonites 3-4, 1 genus 2-4; ab-
domen of single reduced article. Two subfamilies:

Aeginellinae Vassilenko, 1968, emend.

Pereopods 3-4 absent, gills on pereonites 3-4 except 1 genus.
Seven genera: Aeginella, Aeginina, Parvipalpus, Proaeginina,
Protellina, Pseudaeginella, Thorina.

Protellinae new subfamily

Pereopods 3-4 absent or number of articles quite reduced,
gills on pereonites 3-4. Sixteen genera: Abyssicaprella, Deu-
tella, Eupariambus, Liropus, Luconacia, Mayerella, Metapro-
tella, Monoliropus, Orthoprotella, Paraprotella, Parvipalpina,
Protella, Protellopsis, Pseudoprotella, Triliropus, Tritella.

CAPROGAMMARIDAE KUDRJASCHOV AND VASSILENKO,
1966, emend.

Mandible with molar and palp (Group II) or lacking molar
(Group II, aberrant), pereopods 3-4 reduced to | or 2 articles,
gills on pereonites 2-4 or 3-4, abdomen of numerous segments.
Two genera: Caprogammarus, Cercops.

CAPRELLIDAE Wutre, 1847, emend.

Mandible with molar, lacking palp (Group III) except one
genus with rudimentary palp; pereopods 34 quite reduced or
absent; gills on pereonites 3-4; abdomen of single reduced arti-
cle. Nine genera: Caprella, Eugastraulax, Hemiaegina, Meta-
caprella, Paracaprella, Pariambus, Pedoculina, Propodalirius,
Pseudolirius.

840 Proceedings of the Biological Society of Washington

a
~

—

eae AEGINELLINAE

AEGINELLIDAE

STA eet
ee. .° Sk ZO ee
£ x, € Y ;
(4... \ , |
CYAMIDAE \ Ss \ :

PODOCERID-LIKE
ANCESTOR CA PROGAMMARIDAE

yo
PHTISICINAE DODECADINAE C4PRELLIDAE
\ na
\ / ae
Pa Pe fe =

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_
PHTISICIDAE PROTELLINAE 5 |

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GAMMARIDEA

Fic. 1. Familial relationships in the suborder Caprellidea. (Posterior
pereopods omitted except where taxonomically significant)

CYAMDDAE Waite, 1847

Mandible without molar or palp (Group IV), pereopods 34
absent, gills on pereonites 3-4, abdomen of single reduced
article. Five genera: Cyamus, Isocyamus, Neocyamus, Platy-
cyamus, Syncyamus.

Incertae Sedis

Six genera: Aciconula, Noculacia, Paradeutella, Proliropus,
Triantella, Triperopus. Mandibles of these six genera have not
been described fully; however, all have a 3-segmented mandib-
ular palp and appendages on pereonites 3-4. They, therefore,
probably belong to the Protellinae of the Aeginellidae.

Caprellid families 841

Two stocks are apparent within the Protellinae. Four genera
bear reduced fifth pereopods, Eupariambus, Liropus, Mayer-
ella, and Parvipalpina, while in most of the other genera they
are fully segmented. These two stocks may ultimately be
placed in separate subfamilies but their separation at this time
seems unjustified because of the close similarity of their char-
acters.

With the exception of Fallotritella, all Dodecadinae bear
reduced fifth pereopods. The mandible of Fallotritella differs
from that of other Phtisicidae in that the accessory plates are
quite small, almost spinelike. It is possible that Fallotritella
is an example of convergence of the subfamilies Protellinae
and Dodecadinae based on the reduction of the molar on a
typical Group II mandible.

The mandible of Cercops lacks a molar but in other respects
resembles the Group II mandible, lacking the accessory plates
found in the other genera which have no molar. Because of the
segmentation of the abdomen, I have chosen to place it in the
Caprogammaridae.

This system of classification shows several evolutionary
lines within the Caprellidea. Postulating a podoceridlike an-
cestor (McCain, 1968) for the Caprellidea, 3 distinct lines
emerge (Fig. 1). One line gives rise to the Cyamidae with
quite reduced mouthparts, an absence of appendages on pere-
onites 3-4, and bearing gills only on pereonites 3-4. The
Cyamidae are highly specialized parasites which led Barnard
(1969, p. 21) to state that the cyamids comprise a fifth major
group of Amphipoda. Their separation from the Caprellidea
into a fifth suborder deserves consideration by a cyamid
specialist. A second line gives rise to the Phtisicidae in which
all members lack a mandibular molar and most have 3 pairs
of gills. The third line passes through the Caprogammaridae
which have large abdomens and typical mandibles reminiscent
of the podocerids. Farther along this line are the Aeginellidae
with reduced abdomens but typical mandibles. With the loss
of the mandibular palp, the third line ends with the Caprel-
lidae. The genus Paracaprella is intermediate between the
Aeginellidae and the Caprellidae based on its rudimentary palp.

842 Proceedings of the Biological Society of Washington

LITERATURE CITED

BARNARD, J. L. 1969. The families and genera of marine gammaridean
Amphipoda. Bull. United States Nat. Mus., vol. 271, pp.
vi-535, 173 figs.

KupryascHov, V. A. AND S. V. VAssILENKO. 1966. A new family Cap-
rogammaridae (Amphipoda, Gammaridea) found in the
North-West Pacific. Crustaceana, vol. 10, pt. 2, pp. 192-198,
4 figs.

McCaw, J.C. 1968. The Caprellidae (Crustacea: Amphipoda) of the
Western North Atlantic. Bull. United States Nat. Mus., vol.
278, pp. vi-147, 56 figs.

VASSILENKO, S. V. 1968. On the question of systematics and the basic
lines of development in the family Caprellidae [in Russian].
Doklady Akademii Nauk S.S.S.R. [Proc. Acad. Sci. U.S.S.R.],
vol. 183, no. 6, pp. 1461-1464.

Wuire, A. 1847. List of the specimens of Crustacea in the collection of
the British Museum. London. pp. viii-143.

iN

pp. 843-846 5 February 1970

PROCEEDINGS
OF THE

BIOLOGICAL SOCIETY OF WASHINGTON

HATSCHEKIA PACIFICA NEW SPECIES (COPEPODA:
CALIGOIDA) A PARASITE OF THE SAND BASS,
PARALEBRAX NEBULIFER (GIARD)

By Rocer F. Cressry
Smithsonian Institution, Washington, D. C.

As part of a survey of the copepods parasitic on the inshore
fishes of La Jolla, California this paper describes a new species
of Hatschekia contained in six collections from the gills of
Paralebrax nebulifer (Giard ).

All collections were made by Mr. Edmund Hobson and Mr.
Lloyd Richards of the Tiburon Marine Laboratory, U. S. Bu-
reau of Sport Fisheries and Wildlife.

All material has been deposited in the Smithsonian Institu-
tion, Division of Crustacea.

Hatschekia pacifica new species
Figures 1-9

Material studied: Holotype 2 (USNM 126976) and 67 paratype 2 2
(USNM 126977) collected from the gills of Paralebrax nebulifer at
La Jolla, California 2 October 1968. Five additional collections from
the same host and locality were made on 22 July 1968 (2 @ 2? ), 8 August
1968 (2 9 9), 24 September 1968 (5 9 2), 1 October 1968 (3 2 2),
and 2 October 1968 (2 92).

Female: Body form as in figure 1. Total length 2.48 mm. Greatest
width 0.6 mm. Cephalon comprises about one-eighth total length.
Thoracic segment bearing first and second pairs of legs distinct.

Genital segment comprises approximately 75 percent of body length.
Dorsal body surface without ornamentation. Posterior corners of genital
segment smoothly rounded. Abdomen small and one-segmented. Caudal
rami (fig. 2) small, about three times as long as wide, and bearing six
setae, no ornamentation of the rami or setae could be seen under high#e>
magnification. =

First antenna (fig. 3) three-segmented; last two segments incomplethfg
divided, all segments bearing naked setae as in the figure. Second
antenna (fig. 4) in form of a stout claw, penultimate segment covered

66—Proc. Brot. Soc. Wasu., Vou. 82, 1970 (843)

844 Proceedings of the Biological Society of Washington

Hatschekia pacifica new species 845

with fine spinules on distal half. Mandible blade (fig. 5) narrowing
abruptly at distal fourth and terminating as two teeth, entire structure
lying within mouth cone. First and second maxillae (fig. 6) each con-
sisting of two broad but weakly sclerotized setae; first maxilla setae short
and furcalike. Maxilliped (fig. 7) 4-segmented, terminating as a bifid
claw and with setae borne on the inner distal corners of the penultimate
and antipenultimate segments.

First leg (fig. 8) biramose; exopod 2-segmented and armed as in the
figure, endopod 1-segmented and bearing five setae: all segments with
rows of spinules as indicated in the figure. Second leg (fig. 9) biramose;
each ramus 2-segmented and armed as in the figure, rows of spinules on
both exopod segments and the last endopod segment.

Egg strings uniseriate and generally about twice the length of the
body, each string containing 50-75 eggs.

Male: Unknown.

Remarks: This new species differs from most of the known species of
Hatschekia on the basis of the separation of the leg bearing segment from
the cephalon. It seems most closely related to H. conifera Yamaguti but
can be easily distinguished from it because of the processes present on the
terminus of the genital segment of conifera; it can be further separated
by the nature of the armature of the first and second legs, the setae of
pacificus being much longer than in conifera.

This new species is only the third recorded from eastern Pacific
waters. The other two are conifera by Cressey 1968 and pinguis Wilson
1908.

LITERATURE CITED

Cressey, R. F. 1968. A redescription of Hatschekia conifera, Yama-
guti 1939, (Copepoda, Caligoida), including the first de-
scription of the male. Proc. Biol. Soc. Wash., 81 (21): 173-
178.

Witson, C. B. 1908. North American parasitic copepods: a list of
those found upon the fishes of the Pacific coast, with de-
scriptions of new genera and species. Proc. U. S. Nat. Mus.
35 (1652): 431-481.

YamacutTt, S. 1939. Parasitic copepods from fishes of Japan. Pt. 5.
Caligoida, III. Vol. Jub. Prof. S. Yokhida, 2: 443-487.

<

Fics. 1-9. Hatschekia pacifica new species, female: 1, dorsal view;
2, caudal ramus, ventral; 3, first antenna; 4, second antenna; 5, blade of
mandible; 6, first and second maxilla; 7, maxilliped; 8, first leg; 9, second
leg. All drawings except figure 5 drawn with the aid of a camera lucida
—figure 5 freehand.

846 Proceedings of the Biological Society of Washington

No. 67, pp. 847-850 5 February 1970

PROCEEDINGS
OF THE

BIOLOGICAL SOCIETY OF WASHINGTON
a ee BE A

A NOTE ON THE GENERIC NAMES CYCLAGRAS COPE
AND LEJOSOPHIS JAN (REPTILIA: SERPENTES)

By James A. PETErs
Smithsonian Institution, Washington, D. C.

Hoge (1958: 221) recently reviewed the status of the generic
name Lejosophis Jan, 1863, concluding that Dunn (1944: 70)
was correct in using it to replace Cyclagras Cope, 1885. He
presented a synonymy of the monotypic genus, using the name
Lejosophis for it. Hoge’s action was based on the statement
by Dunn (1944: 70) that Boulenger (1894: 144), acting as first
reviser, fixed the type-species of Lejosophis (spelled Leiosophis
by Boulenger) as Xenodon gigas Duméril, and therefore Le-
josophis and Cyclagras became objective synonyms, having the
same type-species. If Dunn’s interpretation is valid, there can
be no question that Lejosophis is the correct name for the
genus, and it would require a petition to the International
Commission of Zoological Nomenclature to set it aside in favor
of the long-established Cyclagras.

Dunn’s action was interpretive in nature. When the British
Museum Catalogues were published by Boulenger, he did not
designate type-species in any manner. He clearly did not like
tautonymic names, and I know of none coined by him in the
Catalogues. A type-species by monotypy in Boulenger’s work
is obvious. A new genus described in his Catalogues and in-
cluding several species must await a subsequent reviser for
designation of a type, because there is never any clue as to his
intent. One occasionally can take advantage of Boulenger’s
style in the Catalogues as a basis for considering an action as
type-species designation, post facto. This, in fact, is what
Dunn has done.

67—Proc. Brot. Soc. Wasu., Vou. 82, 1970 (847)

848 Proceedings of the Biological Society of Washington

Boulenger (1894: 144) gives a synonymy of the genus Cycla-
gras which includes the following lines:

“Xenodon, part., Dum. and Bibr., Erp. Gén. vii. p. 753 (1854).

“Leiosophis, part., Jan, Arch. Zool. Anat. Phys. ii. 1863, p.
320.

“Cyclagras, part., Cope, Proc. Am. Philos. Soc. xxii. 1885,
joe Kota

Since Boulenger then used the junior synonym Cyclagras for
the single species included (Xenodon gigas Duméril), it is
clear that he did not consider Leiosophis available for the
taxon. By using Cyclagras, however, for a monotypic genus,
Boulenger designated gigas as type-species of Cyclagras,
through the exclusion of any other species, or by monotypy.
Since Cope originally described Cyclagras as a substitute name
for Lejosophis, Dunn extended the argument to include Lei-
osophis, and concluded that Boulenger simultaneously desig-
nated gigas as its type-species. Dunn’s reasoning would be
acceptable if this were the only place where Leiosophis was
mentioned, but Boulenger (1894) again referred to the genus
on p. 180, where the pertinent lines read:

“Cosmiosophis, Jan, Arch. Zool. Anat. Phys. ii. 1863, p. 289.

“Leiosophis, part., Jan, l.c. p. 320.”
These citations are in the synonymy of Urotheca Bibron. The
point here is that the two citations to Leiosophis are identical,
and are both referred to as “part.,” or partim, Boulenger’s way
of indicating that only some, not all, of the species assigned
to the genus by the original author are included in the genus
being discussed. Jan included two species in Lejosophis when
he described it, Xenodon gigas Duméril and Coluber bicinctus
Hermann. In Boulenger, the “Leiosophis, part.” under Cycla-
gras refers to gigas, and the “Leiosophis, part.” under Urotheca
refers to bicinctus. It is clearly invalid to say one of these can
be interpreted as a restriction of type-species while the other
is ignored.

It is possible on other grounds to arrive at a type-species for
Leiosophis. As pointed out above, Jan (1863) assigned two
species to his new genus. In the Iconographie Générale des

Notes on Cyclagras and Lejosophis 849

Ophidiens (1881), however, he pointed out in his index that
gigas was to be placed in genus Xenodon and figured it (Livr.
48, pl. 3, fig. 6) under the name Xenodon gigas. Thus, one
could claim that Jan himself has designated bicinctus Hermann
as the type-species of Lejosophis through the same “exclusion
principle” followed by Boulenger in the case of Cyclagras. Jan
vacillated in his treatment of the name gigas, because it was
given as Lejosophis gigas on a different plate (Livr. 50, pl. 2,
figs. 25-27). In his index, under Lejosophis gigas, he wrote
“voy. Xenodon gigas.” Under the genus Xenodon in the index,
he wrote “___ [for Xenodon] (Lejosophis) gigas Dum. Bibr.,”
which makes it look like a subgenus! Jan did not refer to the
second species, Lejosophis bicinctus Hermann, in the Icono-
graphie.

Cope (1885: 185) rejected Lejosophis Jan, indicating that
he felt that Jan had misspelled the name, and that, properly
spelled, it would be Liophis, a preoccupied name (Liophis
Wagler, 1830). Cope mentioned both gigas Duméril and
bicinctus Hermann in his discussion, so he clearly intended to
include both names in his Cyclagras, which he coined as a
replacement name for Lejosophis. No type-species for Cycla-
gras was designated until Boulenger restricted the name to
gigas Duméril, thus fixing that name as the type-species. It is
possible to argue that Cyclagras, since it was proposed as a
replacement name for Lejosophis, and must therefore take the
same generotype, has as its type-species bicinctus Hermann,
and would thus be a generic synonym of Hydrodynastes Fitz-
inger, 1843. This would require a new generic name for gigas
Duméril, an action [ consider unnecessary, superfluous, and a
flouting of the basic concept of stability in zoological names.

If, however, herpetologists are willing to accept the inter-
pretation that Jan removed the taxon gigas from Lejosophis,
transferring it to Xenodon, and thus automatically designated
Coluber bicinctus Hermann as type-species by monotypy, all
problems are resolved. This interpretation will be followed
in the Catalogue of Neotropical Squamata now being prepared
by Braulio Orejas-Miranda and myself. There will be two
genera recognized, as follows:

850 Proceedings of the Biological Society of Washington

Hydrodynastes Fitzinger

1843 Hydrodynastes Fitzinger, Systema Reptilium: 25.
Type-species: Elaps Schrankii Wagler, 1824 (= Col-
uber bicinctus Hermann), by original designation.

1863 Lejosophis Jan, Arch. Zool. Anat. Phys., 2: 320.
Type-species: Coluber bicinctus Hermann, by sub-
sequent monotypy. (See discussion above.)

1944 Dugandia Dunn, Caldasia, 3(11): 70. Type-species:
Coluber bicinctus Hermann, by original designation.

CONTENT: A single species; Hydrodynastes bicinctus
(Hermann).

Cyclagras Cope
1885 Cyclagras Cope, Proc. Amer. Phil. Soc., 22: 185.
Type-species: Xenodon gigas Duméril, by subse-
quent monotypy (in Boulenger, Cat. Sn. Brit. Mus.,
2, 1894, 144).

CONTENT: A single species; Cyclagras gigas Duméril.

LITERATURE CITED

BouLencEr, G. A. 1894. Catalogue of the Snakes in the British Mu-
seum (Natural History), Vol. 2, London, pp. i-xi + 1-382,
20 pls.

Corr, E. D. 1885. Twelfth Contribution to the Herpetology of Trop-
ical America. Proceedings of the American Philosophical
Society, 1884 (1885): 167-194, 1 pl.

Dunn, E. R. 1944. Dugandia, a New Snake Genus for Coluber Bi-
cinctus Hermann. Caldasia, 3(11): 69-70.

Hocr, A. R. 1958. Tres Notas Sobre Serpentes Brasileiras. Papéis
Avulsos de Departmento de Zoologia, So Paulo, Brasil, 13
(7): 221-225.

Jan, G. 1863. Prodromo Della Iconografia General Degli Ofidi, II*
Parte, VI° Gruppo. Coronellidae. Arch. Zool, Anat. Fis., 2
(2): 213-330.
AND F. SorpeLur. 1860-1881. Iconographie Générale des
Ophidiens. Milan and Paris.

GH 7F

nl
La

Va ~ 851-864 5 February 1970

PROCEEDINGS
OF THE

“BIOLOGICAL SOCIETY OF WASHINGTON

NEW ENTOCYTHERID OSTRACODS FROM
TENNESSEE AND VIRGINIA

By Horton H. Hosss, JR. AND MARGARET WALTON
Smithsonian Institution and
Mountain Lake Biological Station

Four new ostracods belonging to the genera Ascetocythere
and Dactylocythere are described from the upper Tennessee
and Cumberland drainage systems in Tennessee and Virginia.

In examining specimens of Dactylocythere spinata (see be-
low), we observed, for the first time, an unpaired, heavily
sclerotized spinelike prominence, here designated the sternal
spine, which extends posteriorly along the ventromedian line
of the body between the first pair of legs. Although this spine
was first observed in Dt. spinata, it occurs, in various forms,
in males of at least 10 of the 27 described members of the genus
(see below) and is particularly well-disposed for illustration
in a paratypic male of Dt. xystroides Hobbs and Walton, 1963:
460, from Hurricane Creek, southeast of Waverly, Humphreys
County, Tennessee (see Figs. la, b).

The sternal spine (ss) appears to articulate anteriorly with
a pair of long, slender, paramedian ventral prongs (vp) which
extend anteriorly between the bases of the maxillae (mx) and
mandibles (md), and to be supported dorsolaterally by the
posteroventral horns (pvh) of paired \-shaped apodemes, the
anteroventral horns (avh) of which are continuous with the
respective right and left ventral prongs. Each of the latter
bears a prominence (mds) which supports the mandible and
from which a slender, simple trabecula extends to the maxilla
of the respective side. The dorsal horn (dh) of the apodeme

bears an anterior spur which supports a complex trabecula ‘:
extending from the proximal base of the maxilla to the base of |

the mandible.
68—Proc. Bron. Soc. Wasu., Vou. 82, 1970 (S59)

‘\
= \
5 \
ae |
Lf.

852 Proceedings of the Biological Society of Washington

Fic. 1. Dactylocythere xystroides. a, Dextral view showing position
of sternal spine (black); b, Apodeme and trabeculae associated with
sternal spine. (See text for explanations of abbreviations. )

Rioja (1940 and 1941) made a careful study of the endo-
skeleton of Ankylocythere heterodonta (Rioja, 1940: 594) [=
Entocythere heterodonta] but did not recognize most of the
elements just described. Not only did we fail to find them in
Rioja’s species but we were also unable to identify them in
certain species of the genus Dactylocythere, and only part of
them were evident in others; thus, it is highly probable that if

New entocytherid ostracods 853

they are present in Ank. heterodonta and in the members ot
the genus Dactylocythere not listed below, they are not strongly
sclerotized and consequently are not visible in available prep-
arations.

Those species of Dactylocythere which possess sternal spines
are: Dt. amicula Hart and Hart, 1966: 1; Dt. brachystrix Hobbs
and Walton, 1966: 2; Dt. chalaza (Hobbs and Walton, 1962:
45); Dt. chelomata (Crawford, 1961: 242); Dt. daphnioides
(Hobbs, 1955: 325); Dt. exoura Hart and Hart, 1966: 5; Dt.
pachysphyrata Hobbs and Walton, 1966: 3; Dt. runki (Hobbs,
1955: 330); Dt. spinata, new species; and Dt. xystroides Hobbs
and Walton, 1963a: 460.

Three of the remaining species of the genus—Dt. jeanae
Hobbs, 1967: 6; Dt. striophylax (Crawford, 1959: 157); and
Dt. suteri (Crawford, 1959: 162)—possess the paired A-shaped
apodemes and ventral prongs, and the posterior extremities of
the latter are produced posteriorly into small lobes, but no
remnant of the sternal spine has been observed in any of the
three.

Those species of the genus which are not listed above appear
not to have the A-shaped apodemes, and the ventral prongs, if
present, are not sclerotized in our specimens.

Acknowledgments: We wish to thank Raymond W. Bou-
chard, Perry C. and Virgie F. Holt for furnishing us with the
specimens on which Ascetocythere holti and Dactylocythere
spinata are based. For criticisms of the manuscript, we are
indebted to Fenner A. Chace, Jr.

Ascetocythere holti new species
(Figures 2a, b, 3a, b)

Male: Eye pigmented. Shell (Fig. 2a) ovate in silhouette but
slightly concave anteroventrally, greatest height slightly posterior to
midlength. Submarginal setae anteriorly, posteriorly, and ventrally; those
situated anteriorly progressively farther from margin dorsally; setae
apparently absent dorsally.

Copulatory complex (Figs. 3a, b) with peniferum bearing three prom-
inences extending from subterminal expansion: anterior process flat-
tened, its length about half that of anteroposterior plane of distal portion
of peniferum, and directed anteroventrally with distal portion deflected
more ventrally; ventral process slightly heavier, subequal in length to ante-

854 Proceedings of the Biological Society of Washington

h

Fic. 2. Right valves of shells. a, d, e, h, Males; b, c, f, g, Females;
a, b, Ascetocythere holti new species; c, d, Dactylocythere enoploholca
new species; e, f, Dactylocythere myura new species; g, h, Dactylocythere
spinata new species.

rior process, directed ventrally, and bearing fold along proximoposterior
margin serving as penis guide; posterior process, situated immediately
posterior to ventral process, slightly undulating, acute, about one-half as
long as latter, and also directed ventrally. Penis complex long and
extending ventrally along penis guides on posterior surface of ventral
process. Clasping apparatus not clearly divisible into vertical and hori-
zontal rami; internal border gently curved between broad base and

New entocytherid ostracods 855

tapering distal portion, and bearing two or three inconspicuous eleva-
tions along distal third (distal elevation acute, almost toothlike in some
specimens ); distal extremity with three anterodorsally directed denticles;
external border also rounded with broadly oblique subangular bend;
extensions of principal proximal and distal axes forming angle of ap-
proximately 105 degrees; height of distal extremity of tapering apparatus
only approximately one-fourth anteroposterior diameter of base of ap-
paratus. Dorsal finger comparatively stout and terminating in bifid
seta extending posteroventrally; ventral finger moderately heavy, dis-
posed somewhat subparallel to anterior margin of ventral portion of peni-
ferum, with one subangular bend, and directed posteroventrally.

Triunguis Female: Eye pigmented. Shell (Fig. 2b) similar in shape to
that of male but distinctly higher in posterior third; submarginal setae
disposed as in male.

Genital complex consisting of sclerotized papilla surrounded by
amorphous hyaline material (presumably the spermatophore) with
dangling, irregularly shaped, delicate saclike membrane.

Measurements (in millimeters ):

Holotype Males Allotype Females
Number of specimens 10 10
Length (range) 0.39: 0.37-0.41 0.40 0.38—0.40
Average 0.39) 0.39
Height (range) 0.22 0.21-0.24 0.24 0.24—0.25
Average 0.22 0.24

Type-locality: Stream, 8.3 miles west of junction of county roads 2451
and 3387 on latter, southeast of Oneida, Scott County, Tennessee.

Disposition of Types: The hclotypic male and allotype are deposited
in the National Museum of Natural History (Smithsonian Institution )
no. 126974. Paratypes are in the collections of C. Willard Hart, Jr. (1 4,
1 9), H. H. Hobbs III (1 ¢,1 2), and in the Smithsonian Institution
(22) 653 9);

Hosts: Cambarus (Depressicambarus) sphenoides Hobbs, an unidenti-
fied crayfish related to Cambarus (Jugicambarus) distans Rhoades, and
another related to Cambarus (J.) obeyensis Hobbs and Shoup.

Range and Entocytherid Associates: TENNESSEE (Cumberland River
drainage system)—Anderson County: ‘Tributary to New River, 4.4
miles E. of Shea, with no entocytherid associates. Campbell County:
Small stream, 1.5 miles E. of Shea, with Donnaldsoncythere tuberosa
(Hart and Hobbs, 1961: 182) and Dactylocythere sp.; Small stream,
9 miles S.W. of Caryville on road to Shea, with Dn. tuberosa, Dt. spinata,
and Entocythere sp. Fentress County: Campbell Branch, 0.4 mile N.W.
of junction of Tenn. Rte. 52 and unnumbered road near Armathwaite,

856 Proceedings of the Biological Society of Washington

New entocytherid ostracods 857

with Dn. tuberosa and Dactylocythere sp. Morgan County: Mud Creek
at Tenn. Rte. 52, with Dn. tuberosa; White Oak Creek on U.S. Hwy. 27
at Sunbright, with Dn. tuberosa and Dt. spinata. Scott County: Type-
locality, with Dn. tuberosa and Dt. sp.; Bandy Creek W. of Leather-
wood Fork, with Dn. tuberosa, Dt. spinata, and Entocythere sp.; Painted
Rock Creek on Tenn. Rte. 63, E. of Huntsville, with Dn. tuberosa;
Perkins Creek at U. S. Hwy. 27, N.E. of Winfield, with Dn. tuberosa,
Dactylocythere sp., and Entocythere sp.

Relationships: Ascetocythere holti is a member of the Asceta Group
of the genus and seems to have its closest affinities with A. sclera Hobbs
and Hart, 1966: 42. It shares with all of the species of the group a
clasping apparatus in which the major bend occurs proximal to its
midlength, and with A. sclera, A. didactylata Hobbs and Hart, 1966: 43,
and A. batchi Hobbs and Walton, 1968: 237, the absence of a flangelike
process projecting from the ventral surface of the peniferum. It differs
from A. didactylata in possessing three processes on the ventral portion
of the peniferum, from A. sclera in having a much longer anterior process,
an acute, undulating posterior process, and a subangular bend on the
posteroventral margin of the peniferum, and from A. batchi in possessing
a posterior process.

Etymology: It is a pleasure to name this species in honor of our good
friend and colleague, Dr. Perry C. Holt, who has contributed numerous
specimens of crayfishes and entocytherids to us.

Dactylocythere enoploholea new species
(Figures 2c, d, 3c, d)

Male: Eye pigmented, situated approximately one-eighth shell length
from anterior margin. Shell (Fig. 2d) ovate with greatest height dis-
tinctly posterior to midlength. Submarginal setae present except dorsally
between level of posterior margin of eye and that of dorsal portion of
peniferum. Sternal spine lacking.

Copulatory complex (Figs. 3c, d) with finger guard broad at base and
tapering to form narrow distal portion, ventral margin emarginate with
posteriorly directed acute tip; peniferum moderately heavy with sub-
truncate ventral margin and posteroventral “heel”; accessory groove
reaching level slightly dorsal to dorsal margin of spermatic loop, with
its dorsalmost portion folded and possessing irregular margin; peniferal
groove opening anteriorly, its apical width approximately two-thirds that

<

Fic. 3. Copulatory complexes. a, c, f, h, Entire complexes drawn to
scale 1; b, d, e, g, Finger guards and clasping apparati drawn to scale 2;
a, b, Ascetocythere holti new species; c, d, Dactylocythere enoploholca
new species; e, f, Dactylocythere myura new species; g, h, Dactylocythere
spinata new species.

858 Proceedings of the Biological Society of Washington

of diameter of vertical ramus of clasping apparatus above rounded
shoulder (see below); penis somewhat L-shaped, situated in ventral
fourth of peniferum; clasping apparatus, extending ventrally beyond
ventral margin of peniferum, with two major bends, but not clearly
divisible into vertical and horizontal rami and with major axes of ex-
tremities forming angle of approximately 70 degrees; external borders of
both rami entire but that of vertical ramus with rounded shoulder at
midlength (level of proximal bend); internal border of horizontal ramus
with one large tooth near midlength and with three low elevations
immediately proximal to three apical denticles. Both dorsal and ven-
tral fingers moderately slender, latter more than twice length of former,
gently curved from base and suddenly curved posteriorly at base of
distal third.

Triunguis Female: Eye pigmented and situated as in male. Shell
(Fig. 2c) distinctly larger than that of most males, more highly vaulted
posteriorly, and with shallow ventral excavation anterior to midlength.
Submarginal setae present except dorsally between level of eye and
genital complex.

Genital complex consisting of prominent, but short, J-shaped rod and
amiculum, latter protruding little, if at all, between valves.

Measurements (in millimeters ):

Holotype Males Allotype Females
Number 10 10
Length (range) 0.46 0.44—0.49 0.47 0.46—0.55
Average 0.45 0.50
Height (range) 0.25 0.25-0.28 0.29 0.28-0.34
Average 0.27 0.30:

Type-locality: South Fork of the Holston River at junction of state
routes 600 and 762, Washington County, Virginia. This is the only
locality in which this species is known to occur.

Disposition of Types: The holotypic male and allotype are deposited
in the National Museum of Natural History (Smithsonian Institution )
no. 126973. Paratypes are in the collections of C. Willard Hart, Jr. (1 ¢,
1 @), H. H. Hobbs III (1 6,1 @), and in the Smithsonian Institution
(3 6,39).

Hosts: The type-series was obtained from a collection of Cambarus
(Hiaticambarus) longirostris Faxon and Cambarus (Puncticambarus )
sp.

Entocytherid Associate: Dactylocythere falcata (Hobbs and Walton,
1961: 379).

Relationships: Dactylocythere enoploholca has as its closest relatives
Dt. chalaza, Dt. pachysphyrata, and Dt. spinata. In all four species, the
accessory groove extends dorsally approximately to the level of the

New entocytherid ostracods 859

spermatic loop; the aperture of the peniferal groove is directed ante-
riorly; and the clasping apparatus is curved at almost the same angle
and bears only one major tooth (two in Dt. pachysphyrata) on the in-
ternal border of the clasping apparatus. Dactylocythere enoploholca
differs from the other three, however, in having a heel-like prominence
on the posteroventral margin of the peniferum, a prominent rounded
shoulder on the external border of the vertical ramus of the clasping
apparatus, and in lacking a sternal spine.

Etymology: Enoplus (Greek) = armed, and holkos = furrow; so
named because of the folded irregular dorsal extremity of the accessory
groove of the peniferum of the male.

Dactylocythere myura new species
(Figures 2e, f, 3e, f)

Male: Eye pigmented, situated approximately one-fifth shell length
from anterior margin. Shell (Fig. 2e) elongate ovate with greatest height
at midlength. Submarginal setae present anteriorly, posteriorly, and
ventrally, but none present dorsally between level of eye and dorsal
portion of peniferum. Sternal spine lacking.

Copulatory complex (Figs. 3e, f) with finger guard rather heavy, its
posterior margin concave and its oblique distal margin with three
prominences of which anteriormost extending considerably farther ven-
trally than posterior one; peniferum moderately heavy with rounded
ventral margin lacking tubercles, emarginations, or scallops; accessory
groove reaching level of dorsal margin of spermatic loop with simple
round dorsal extremity; peniferal groove opening anteriorly, its apical
width narrow, no more than one-fourth least diameter of vertical ramus of
clasping apparatus; penis L-shaped and situated at base of distal fourth
of peniferum; clasping apparatus also L-shaped with vertical ramus
slightly bowed anteriorly, extending ventrally beyond peniferum, clearly
divisible into vertical and horizontal rami, and major axes forming angle
of approximately 85 degrees; external borders of both rami and internal
border of vertical ramus entire, that of vertical ramus lacking shoulder; in-
ternal border of horizontal ramus usually without teeth but with three low
elevations distal to midlength (elevations occasionally subacute), and
bearing three or four small dorsally directed denticles. Dorsal finger
somewhat heavier than ventral and terminating in bifid tip; ventral
finger gently curved throughout its length.

Triunguis Female: Eye pigmented and situated slightly more anteriorly
than that of male. Shell (Fig. 2f), while scarcely longer than that of
male, distinctly more highly vaulted with greatest height posterior to
midlength, and with much steeper slope posterodorsally; ventral margin
with only faintest indication of shallow excavation anterior to midlength.
Submarginal setae distributed as in male.

Genital complex consisting of prominent, long J-shaped rod and long
amiculum, latter sometimes slightly protruding between valves.

860 Proceedings of the Biological Society of Washington

Measurements (in millimeters):

Holotype Males Allotype Females
Number 9 5
Length (range) 0.48 0.46-0.48 0.48 0.48
Average 0.47 0.48
Height (range) 0.27 0.25-0.27 0.29 0.27-0.29

Average 0.27 0.28

Type-locality: Burrows along bank of spring-fed stream, 3.5 miles
southwest of Chilhowie in Washington County, Virginia (Holston River
drainage system).

Disposition of Types: The holotypic male and allotype are deposited
in the National Museum of Natural History (Smithsonian Institution )
no. 126975. Paratypes are in the collections of C. Willard Hart, Jr. (1 ¢,
2 9), H. H. Hobbs III (1 6, 1 @), and in the Smithsonian Institution
(4 6,2 9).

Host: An undescribed crayfish closely allied to Cambarus carolinus
Erichson.

Range and Entocytherid Associates: Dactylocythere myura is known
from only one locality other than the type-locality, 1.0 mile southwest
of Chilhowie off Interstate Hwy. 81, Smyth County, Virginia—only 2.5
miles from the type-locality and also in the Holston drainage system. In
the type-locality, it was associated with Donnaldsoncythere scalis Hobbs
and Walton, 1963b: 364, and in the Smyth County locality with Dn.
scalis and Ascetocythere hyperoche Hobbs and Hart, 1966: 41.

Relationships: While Dt. myura is not obviously closely allied to any
other species of the genus, the comparatively slender clasping apparatus
and the finger guard with three lobes of which the anterior one extends
farthest distally are somewhat like those found in Dt. suteri. It may be
readily separated from the latter, however, by the more angular clasping
apparatus which bears no more than one distinct tooth proximal to the
apical denticles, and the distal margin of the finger guard is oblique
rather than subtruncate. Its more distant relatives include Dt. jeanae
and Dt. striophylax; in neither of these, however, is the finger guard
distinctly trilobed distally.

Etymology: Myurus (Greek) = narrow; alluding to the narrow hori-
zontal ramus of the clasping apparatus of the male.

Dactylocythere spinata new species
(Figures 2g, h, 3g, h)

Male: Eye pigmented, situated slightly more than one-fourth shell
length from anterior margin. Shell (Fig. 2h) elongate ovate with greatest
height some distance posterior to midlength. Marginal setae present
anteriorly, posteriorly, and ventrally. Sternal spine prominent, long,
directed posteriorly with apical portion only slightly bent ventrally.

New entocytherid ostracods 861

Copulatory complex (Figs. 3g, h) with finger guard rather heavy,
its posterior margin deeply convex anteriorly, and its ventral border
excavate with anteroventral prominence decidedly smaller than bituber-
culate, posteroventrally directed posterior prominence, thus ventral mar-
gin with three prominences; peniferum moderately heavy with posterior
margin slightly undulating, but nowhere angulate or with lobes, and
terminating in anteriorly directed acute tip; peniferal groove, only slightly
wider at apex than one-half least diameter of vertical ramus of clasping
apparatus, and directed anteriorly; penis L-shaped and situated approx-
imately at base of ventral fourth of peniferum; clasping apparatus clearly
divisible into vertical and horizontal rami with major axes forming angle
of approximately 80 degrees although vertical ramus with proximal por-
tion bent anteriorly, almost paralleling horizontal ramus. External border
of both rami and internal border of vertical ramus entire and external
border of vertical ramus without conspicuous shoulder; internal border
of horizontal ramus with single large tooth near midlength and two or
three exceedingly low prominences immediately proximal to three small
apical denticles; clasping apparatus thickest in region of junction of two
rami, tapering slightly proximally and distally; portion of horizontal
ramus distal to major tooth not nearly so thick as that proximal to it.

Triunguis Female: Eye pigmented and located slightly more anteriorly
than that of male. Shell (Fig. 2g) much more highly vaulted posteriorly
than in male, with greatest height some distance posterior to midlength
and with posterior margin subtruncate; ventral margin with shallow
excavation at about midlength. Submarginal setae disposed as in male.

Genital complex consisting of prominent J-shaped rod and ruffled
amiculum, frequently with small portion of latter slightly protruding
posteriorly beyond margins of valves.

Measurements (in millimeters ) :

Holotype Males Allotype Females
Number 8 9
Length (range) 0.46: 0.44-0.48 0.48 0.45—0.49
Average 0.46 0.47
Height (range) 0.27 0.27-0.29: 0.30 0.28—0.32
Average 0.27 0.30

Type-locality: Small stream, 9.0 miles southwest of Caryville on
county road to Shea, Campbell County, Tennessee (Cumberland River
drainage system).

Disposition of Types: The holotypic male and allotype are deposited
in the National Museum of Natural History (Smithsonian Institution),
no. 126972. Paratypes are in the collections of C. Willard Hart, Jr. (1 ¢,
1 @), H. H. Hobbs III (1 ¢, 1 2) and in the Smithsonian Institution
(73,62).

862 Proceedings of the Biological Society of Washington

Host: <A crayfish tentatively identified as Cambarus (J.) distans
Rhoades.

Range and Entocytherid Associates: TENNESSEE (Cumberland
River drainage system)—Campbell County: Type-locality, with As.
hoilti, Donnaldsoncythere tuberosa, and Entocythere sp. Fentress County:
Laurel Fork, 9.7 miles N.E. of Jamestown on Tenn. Rte. 154, with Dacty-
locythere sp. Morgan County: White Oak Creek on U. S. Hwy. 27 at
Sunbright, with As. holti and Dn. tuberosa. Scott County: Bandy Creek
W. of Leatherwood Fork, with As. holti, Dn. tuberosa, and Entocythere
sp.
Relationships: Dactylocythere spinata has its closest affinities with
Dt. chalaza and its allies (see discussion of relationships of Dt. enopio-
holca above ), and it is more similar to this species than to the other two.
It differs from Dt. pachysphyrata in having only a single major tooth
on the internal border of the horizontal ramus of the clasping apparatus,
a bituberculate posteroventral prominence on the finger guard, and a
sternal spine that is directed posteriorly rather than ventrally. In Dt.
chalaza, the posteroventral prominence on the finger guard is not
bituberculate, the guard is not strongly bowed anteriorly, and the sternal
spine is directed posteroventrally rather than posteriorly. It differs from
Dt. enoploholca in possessing a sternal spine and in lacking a heel-like
prominence on the posteroventral margin of the peniferum.

Etymology: Spina (L.) = spine; referring to the long sternal spine of
the male.

LITERATURE CITED
CrAawForpD, Epwarp A., Jr. 1959. Five new ostracods of the genus
Entocythere (Ostracoda, Cytheridae) from South Carolina.
Univ. South Carolina Publ., Ser. III, Biol., 2 (4): 149-189,
5 pls.

. 1961. Three new species of the genus Entocythere (Ostra-
coda, Cytheridae) from North and South Carolina. Amer.
Midl. Nat., 65 (1): 236-245, 21 figs.

Hart, C. Witiarp, Jr., AND DasBnry G. Hart. 1966. Four new ento-
cytherid ostracods from Kentucky, with notes on the troglo-
bitic Sagittocythere barri. Notulae Naturae, (338): 1-10, 13
figs.

, AND Horton H. Hosps, Jr. 1961. Eight new troglobitic
ostracods of the genus Entocythere (Crustacea, Ostracoda )
from the eastern United States. Proc. Acad. Nat. Sci.,
Philad., 113 (8): 173-185, 32 figs.

Hosss, Horton H., Jr. 1955. Ostracods of the genus Entocythere
from the New River System in North Carolina, Virginia, and
West Virginia. Trans. Amer. Micros. Soc., 74 (4): 325-333,
10 figs.
1967. A new genus and three new species of ostracods with

New entocytherid ostracods 863

a key to genus Dactylocythere (Ostracoda: Entocytheridae).
Proc. U. S. Nat. Mus., 122 (3587): 1-9, 1 fig.

, AND C. Wiiiarp Hart, Jr. 1966. On the entocytherid
genera Ascetocythere, Plectocythere, Phymocythere (gen.
nov.), and Cymocythere, with descriptions of new species.
Proc. Acad. Nat. Sci., 118 (2): 35-61, 37 figs.

, AND MarcarET WALTON. 1961. Additional new ostracods
from the Hiwassee drainage system in Georgia, North Caro-
lina, and Tennessee. Trans. Amer. Micros. Soc., 80 (4):
379-384, 8 figs.

1962. New ostracods of the genus Entocythere from the
Mountain Lake region, Virginia (Ostracoda Entocytheridae ).
Virginia Journ. Sci., 13 (2): 42-48, 12 figs.

1963a. Three new ostracods (Ostracoda, Entocytheridae )
from the Duck River drainage in Tennessee. Amer. Mid].
Nat., 69 (2): 456-461, 10 figs.

1963b. Four new species of the genus Donnaldsoncythere
(Ostracoda, Entocytheridae) from Virginia with a key to
the species of the genus. Trans. Amer. Micros. Soc., 82
(4): 363-370, 26 figs.

1966. A new genus and six new species of entocytherid
ostracods (Ostracoda, Entocytheridae). Proc. U. S. Nat.
Mus., 119 (3542): 1-12, 2 figs.

1968. New entocytherid ostracods from the southern United
States. Proc. Acad. Nat. Sci., Philad., 120 (6): 237-252,
3 figs.

Rioja, EnriQueE. 1940. Estudios Carcinologicos V. Morfologia de un

ostracodo epizoario observado sobre Cambarus (Cambarellus)
montezumae Sauss. de México, Entocythere heterodonta n.
sp. y descripcién de algunos de sus estados larvarios. An.
Inst. Biol. de la Univ. Nal. A. de México, 11 (2): 593-609,
3 pls.
1941. Estudios Carcinologicos VI. Estudio morfologico del
esqueleto interno de apodemas quitinoso de Entocythere
heterodonta Rioja (Crust. Ostracoda). An. Inst. Biol. de la
Univ. Nal. A. de México, 12 (1): 177-191, 2 pls.

864 Proceedings of the Biological Society of Washington

a Vie

865-870 5 February 1970

PROCEEDINGS
OF THE

DIULUGICAL SOCIETY OF WASHINGTON

NOTES ON THE WEST AMERICAN NEPHROPIDEAN
LOBSTER, NEPHROPSIS OCCIDENTALIS FAXON

By RayMonp B. MANNING
Smithsonian Institution, Washington, D. C.

This is the second in a series of planned reports on nephropid
lobsters, with present emphasis on American species. In the
first paper a striking new genus and species, Nephropides
caribaea, was described from the Caribbean Sea (Manning,
1969). This report includes observations on the only West
American lobster, Nephropsis occidentalis Faxon.

There has been a recent renewal of interest in the biology
and systematics of marine nephropidean lobsters, particu-
larly by those concerned with Indo-West Pacific species (Berry,
1969; Bruce, 1965, 1966a, 1966b, 1966c; Holthuis, 1964; and
Yaldwyn, 1954). As pointed out by several students of the
group, one aspect of the present interest is the availability in
commercial quantities of some nephropids, especially species
of the genus Nephrops.

A survey of American nephropids in the collections of the
Division of Crustacea, National Museum of Natural History,
Smithsonian Institution (USNM), revealed the presence of
several lots of N. occidentalis from unrecorded localities. As
far as I can determine, there are only three records of this spe-
cies in the literature, the original description by Faxon (1893),
which was supplemented by Faxon in 1895, and the extension
of range from the west coast of Mexico to Chile by Bahamonde _
(1959). I take this opportunity to illustrate the pleopod and /= ~ ;
thoracic sternal region of a male. Preliminary studies indicate [= > wy)
that these may provide important characters in the nephropids.'3 © :

I thank Roger F. Cressey for comments on the manuscript. \==

{= N tj

The illustrations are by my wife Lilly. The support of the ‘3

69—Proc. Brot. Soc. WasH., Vou. 82, 1970 (865)

866 Proceedings of the Biological Society of Washington

Nephropsis occidentalis Faxon 867

Smithsonian through its Research Awards Program is acknowl-
edged.

Nephropsis occidentalis Faxon, 1893
(Figures 1-3)

Nephropsis occidentalis Faxon, 1893, p. 195; 1895, p. 127, pl. D, figs. 1
[color], la, 1b.—De Man, 1916, p. 97 [listed, table] Bouvier, 1917,
p. 20 [key].—Balss, 1927, p. 24 [table]_—Bahamonde, 1959, p. 224,
figs. 1-4.

Material: 124, 103 mm; off west coast of Baja California, Mexico;
27°38’45”"N, 115°17'40”W; 525 fathoms; green mud, Globigerina; “Alba-
tross” Station D 5688; 23 April 1911.—19, 101 mm; between Ballenas
Bay and Santa Maria Bay, west coast of Baja California, Mexico; 25°31’
15’N, 113°29'30”’W; 645 fathoms; green mud, fine sand, Globigerina;
“Albatross” Station D 5676; 17 March 1911.—1¢, 108 mm; off Cape
San Lucas, Baja California, Mexico; 22°56’45”N, 109°50/15”W; 630
fathoms; coarse sand, green mud, gravel; “Albatross” Station D 5683;
20 April 1911.—2¢, 101-132 mm; near Trés Marias Islands, Mexico;
21°15'N, 106°23’W; 676 fathoms; gray sand, broken specks; “Albatross”
Station 3424; 18 April 1891; syntypes; USNM 21082.—11 ¢, 51-113 mm;
169, 68-127 mm; off Acapulco, Mexico; 16°33’N, 99°52’30”"W; 660
fathoms; brown sand, broken specks; “Albatross” Station 3418; 11 April
1891; syntypes; USNM 21081.—1¢, 81 mm; off Valparaiso, Chile; ca.
33°S; more than 300 fathoms; John Manning, collector.

Remarks: Relatively little can be added to Faxon’s brief but excellent
original description. The smaller specimens are less pubescent than the
larger ones, and, in smaller specimens, the tubercles on the carapace,
particularly those extending posteriorly from the rostrum, are compara-
tively sharper. The single Chilean specimen shows no marked differences
when compared with Mexican specimens of the same size; however, the
Chilean specimen apparently lacks a middorsal patch of small tubercles
near the posterior border of the carapace which is visible in all of the
Mexican specimens. Faxon (1895) commented on the inflated carapace
in this species. The inflation of the branchial regions is particularly well
marked in specimens longer than 100: mm.

Nephropsis occidentalis resembles five other species in the genus in
having a middorsal carina on the second to fifth abdominal somites. It
further resembles N. aculeata Smith, N. carpenteri Wood-Mason, and
N. rosea Bate in having but one pair of lateral rostral spines. Of the
other species with the middorsal carina on the abdomen, N. ensirostris
Alcock lacks lateral rostral spines and N. atlantica Norman has two pairs.

<

Fics. 1-3. Nephropsis occidentalis Faxon, male, 108 mm, “Alba-
tross” Station D 5683: 1, ventral surface of thorax; 2, male pleopod in
lateral view; 3, male pleopod in mesial view.

868 Proceedings of the Biological Society of Washington

Nephropsis occidentalis differs from all species now known in the genus
in having an erect dorsal spine on the telson near the anterior margin.

Of the American species of Nephropsis, N. occidentalis rather closely
resembles N. aculeata; other than the dorsal spine on the telson and the
projections at the bases of the walking legs discussed below, the two
species are very similar. In N. aculeata the abdominal pleura are nar-
rower and sharper, the chelae are more pubescent, and the body pubes-
cence is not so well-developed.

Examination of the thoracic sternum of males of N. occidentalis
revealed the presence of characters which may prove to be distinctive in
members of the genus. The sternum of a male, 198 mm long, is shown
in Figure 1. On the inner surface of the basal segment of the third
pereiopod there is a spinous triangular projection which is recurved
posteriorly. A similar, sharper projection at the base of the fourth leg is
directed anteriorly. The process on the third pereiopod is directed
ventrally, only slightly recurved, with smooth margins, in males 70 mm
long. In males 90 mm long the apex is recurved posterolaterally, and
at 108 to 113 mm the inner margin is tuberculate. The tubercles are
comparatively much larger than those illustrated in the largest (132. mm)
male examined. In N. aculeata this process on the third leg is larger
than in N. occidentalis and the apex is subdivided into three or four
prominent, sharp, posteriorly directed spines. This character needs to be
surveyed throughout the genus.

Although the male pleopod (Figures 2, 3) has not been used as a
specific character in this group I have included the illustrations here for
future reference.

Nephropsis occidentalis is now known from localities off Mexico be-
tween western Baja California and Acapulco, and from off Chile, where
it was first recorded by Bahamonde (1959). It has not yet been taken
in the Panamanian region, but its absence there may reflect collecting
effort rather than actual occurrence of the species. It apparently occurs
on soft bottom in depths between 525 and 676 fathoms.

In his key to the species of Nephropsis, Bouvier (1917, p. 20) indicated
that N. occidentalis was known from the Galapagos Islands and Iles
Marion; these must be lapsi for Faxon’s original records, both of which
were from off Mexico.

LITERATURE CITED

BAHAMONDE, N. 1959. Decapodos Chilenos: La Familia Homaridae.
Inv. Zool. Chilenas, 5, pp. 221-227, figs. 1-4.

Bauss, H. 1927. Macrura der Deutschen Tiefsee-Expedition. 3. Na-
tantia, Teil B. Wiss. Ergebn. Valdivia Exped., vol. 23,
pp. 247-275, figs. 1-32, pl. 6.

Berry, P. F. 1969. The biology of Nephrops andamanicus Wood-
Mason (Decapoda, Reptantia). South African Assoc. Mar.
Biol. Res., Oceanogr. Res. Inst., Invest. Rep. No. 22, pp.
1-55, figs. 1-26.

Nephropsis occidentalis Faxon 869

Bouvier, E. L. 1917. Crustacés décapodes (macroures marcheurs )

provenant des campagnes des yachts Hirondelle et Princesse-
Alice (1885-1915). Rés. Camp. Sci. Monaco, vol. 50, pp. 1-
140, pls. 1-11.

Bruce, A. J. 1965. On a new species of Nephrops (Decapoda, Rep-

tantia) from the South China Sea. Crustaceana, vol. 9, pt. 3,
pp. 274-284, pls. 13-15.

1966a. Distribution of the genus Nephrops (Crustacea
Decapoda Macrura) in the Indo-Pacific region. Nature, vol.
209, no. 5022, p. 535.

1966b. Nephrops sinensis sp. nov., a new species of lobster
from the South China Sea. Crustaceana, vol. 10, pt. 2, pp.
155-166, pls. 10-12.

1966c. Nephrops australiensis sp. nov., a new species of
lobster from northern Australia (Decapoda Reptantia).
Crustaceana, vol. 10, pt. 3, pp. 244-258, pls. 25-27.

Faxon, Water. 1893. Preliminary descriptions of new species of

Ho.rutuis,

Man, J. G.

Crustacea. Reports on the dredging operations off the west
coast of Central America to the Galapagos, to the west coast
of Mexico, and in the Gulf of California, in charge of Alex-
ander Agassiz, carried on by the U. S. Fish Commission
Steamer “Albatross,” during 1891, Lieut. Z. L. Tanner, U.S.N.,
commanding. VI. Bull. Mus. Comp. Zool. Harvard, vol. 24,
no. 7, pp. 149-220.

1895. The stalk-eyed Crustacea. Reports on an exploration
off the west coasts of Mexico, Central and South America, and
off the Galapagos Islands, in charge of Alexander Agassiz,
by the U. S. Fish Commission Steamer “Albatross,” during
1891, Lieut. Commander Z. L. Tanner, U.S.N., com-
manding. XV. Mem. Mus. Comp. Zool. Harvard, vol. 18,
pp. 1-292, figs. 1-6, pls. A-K, 1-57.

L. B. 1964. On some species of the genus Nephrops
(Crustacea, Decapoda). Zool. Meded. Leiden, vol. 39, pp.
71-78, fig. 1.

DE. 1916. Families Eryonidae, Palinuridae, Scyllaridae and
Nephropsidae. The Decapoda of the Siboga Expedition, Part
III. Siboga Exped. Monogr. 39 (a) (2), pp. 1-122, pls. 1-4.

MANNING, RAyMonp B. 1969. A new genus and species of lobster

(Decapoda, Nephropidae) from the Caribbean Sea. Crus-
taceana, vol. 17, no. 3, pp. 303-309, fig. 1, pl. 1.

YALpwin, J. C. 1954. Nephrops challengeri Balss, 1914 (Crustacea,

Decapoda, Reptantia) from New Zealand and Chatham
Island Waters. Trans. Roy. Soc. New Zealand, vol. 82, no.
3, pp. 721-732, figs. 1-2.

870 Proceedings of the Biological Society of Washington

"

7“

INDEX TO NEW TAXA

VOLUME 82

(New taxa indicated in boldface; n.c. = new combination)

MONOCOTYLEDON

PANDANALES

Pandanus decus-montium — 441
PLATYHELMINTHES

TURBELLARIA
Planaria dactyligera musculosa 543
(iyo rE fie coun ee pe ea er ee a Ee a Ce OBO 549

TREMATODA
MACS GOV TIS. WVeNE 226 es 462
GUISSEVSUre Win eh pe ee 171
amacleithrium — i,

ASCHELMINTHES

KINORHYNCHA
Neocentrophyidae __..---- 116
Neocentrophyes ___.._---- nnn 117
ATULORUYICC RIS © ee 117
SS ER BAY RM pe eae rn, ta lett Fh s 121

NEMATODA

Rhaptothyreidae _- 82
Rhaptothyreus ___.- 82
EY A CC MIS ee a ae 82
S yTriTh ST OTOTINUS) ee ee ee 511
EY, DTC US ae 512

ANNELIDA

POLYCHAETA
Aonides mayaguezensis __.- 393
Apoprionospio _ 383
GASP CTS INTO: C reper Camere eS m neta e e ee eee 381
Cte cet ae A ae PC 383
LOS ETD G Speman a eee Pak NTN een OE EPA RNS hn A 381
DY CUM AC ANTI Ci meter eek Si 381
Cella Fea Yas Woh 01 chine nese POO ees alse aoe ele ees Pane er 381

(871)

872 Proceedings of the Biological Society of Washington

Australaugeneria nnn 20
michaeélseni. 2 ee 22,
PUbIANS NC) oe 22)
pottel New NANG 2 ee 521
Grubeopolynoe _______--- 56
SCMECROVI NG) = eee 59
ETHER STD Ck sc eek eee ee 56
Hololepidella venosa n.c. ___ 22 nnn 50
Neohololepidella — 50
WONTAR oe ee 52
Derren yt: aa a 12
CTINOIGIOO: iy, oo eee 13
Peribaci lates NG. 5 ce 16
Parahololepidella 54
OTOOH TG) accede 5 54
Parapnonospio pinnate ItiCs see 389
Polvewmoa tlyniit 0, oagce a 48
Pottsiscalisetosus 16
PMCIONGNS WHC) <..e e 19
Sthenelanella ehlersi nec. 22 ee 434
LLL | 4 | nn aoe: eee Mmmm enn NNT WEST Cc 8
MIOCDSTOT Ce oe 12
PapUliterd ThC, ee Oe ee 10
DOUUMAA 1.0). xo 8
W@nOnIa. 22.233 ee eee 205
kites pensis: 22. ee eee 205
MOLLUSCA
LAMELLIBRANCHIA
Eupera haitiensis i el ee Be = 825
CEPHALOPODA
Ilex oxygonittis 2 eee 299
ARTHROPODA
CRUSTACEA
Arcania sagamiensis — 247
Ascetocythere holti) 0. 853
Bathyconchoecia deevyae _ 403
Benthochascon elongatum _ nn 259
Bomolochus longieaudus __ 412
Prolixis 220 ee eee 418
spimulus — nnn 420
Cambarus (Cambarus) howardi — 281
Cambarus (Depressicambarus) unestami — 287

Cancer madaensis ______ nnn 258

Capella green eye a
Garcioplax tomentosa:
Chlorinoides tosaensis _.....___._____
Chorisquilla:
Cry ptodnomia: eristatipes 22
Dactylocythere enoploholea —

PET d VE: eae a sa ere Oe

SDD eh ek ee eee i RE ce Bie a ce
TES Cy an en ee ee ee
easel isa Coy i sy 016) Sa nee ae = Rr
Pallicambarus hortoni 22...
Gonodactylopsis. . = ee
TEN capo tS gun MM ex se

Seliher sere te om een enamel
iatschelcnrpaciica: 28.2. ee ol fae Dees ee
Heteropilumnus mikawaensis
lobbselis' attenulatus. 2
Hypolobocera (Phyllothelphusa) niceforoi
Idotea (Pentidotea) kirchanskii
Lepeophtheirus paulus
Leptomithrax kiiensis

Leucosia mimasensis
Macrophthalmus (Macrophthalmus) ceratophorus
Maja nagashimaensis

Neoliomera acutidens

PICHON Oi es) ee
Neorhynchoplax ariakensis __..-_---------__------ en ---
(Ormithocyuiere. SypOdes: sc
@rthotheres:

RE UETE by Ce aaa NY, FF se
Parthenope (Pseudolambrus) ozakii —
Planopilumnus minabensis
Procambarus elegans ___....--__=_ pe

Fg LCD 0) Pay Rescate ee PS
Teo EY 8 Gye a ee
ASO Pole Li 1 cd GS eo se a

SAN PUINGUS: 2 ee
SALE UEC 10s 152 el eR ag A ee
SATITOCY UIVGT G2 ee

CED E98 eaten ee ase ee ee Se
Sphenocarcinus: bidens 8220 2
Mibalarnitacyorunensis. 2 Se ee.
FRAC TLASECOLOS een nm cele Sk eee

ealiforniensis

874 Proceedings of the Biological Society of Washington

DIPLoPopA
Tephlobolelis (2 ee
whiteheadi ____-- ne

CHILOPODA
Cryptops paradrus —

SIPUNCULIDA
ASPIGOSIPHON WINNT 2 oe
Golfingia constricticervix
WVU a eee
murinae bilobata
murinae unilobata _
Onchnesoma magnibatha
ECHINODERMATA

ASTEROIDEA
Litonotaster africanus

CHORDATA

PISCES
Githanchthys “abbott. 22.
Meiacanthus nigrolineatus
Notropis' xanthicara, 2202 eee
Paraclinus fehilmanni 202
Paragunnellichthys fehlmanni —
Splioeroidies ratwins oo
REPTILIA
Agkistrodon contortrix phaeogaster
DISCIVOTUS CONnanh 2 eee
Diploslossus warréni. 20.2 ee
Oligodon analepticos new name ~......-_-.--------------------
AVES

Glyphorhynchus spirurus pallidulus __---------.—------------------------
Habia fuscicauda willist 2.2. eee
Metallura primolinus recisa _____.--__-------_-------------
Oryzoborus crassirostus lottinn se
Sittasomus griseicapillus emochrus
Thamnophilus doliatus nesiotes
Xenops rutilans incomptus

MAMMALIA

Canis adustutus namrui

oe

ES SMITHSONIAN

= -
2 (a
“> | =
Sa) Fes >
J i” ed
/ wn
m
= w
SNI an
z
< =
Paes =
ro) =
mp) 7)
A ‘oS!
- =
>
oi S
ES SMITHSONIAN
wn z
ud 77)
oe ae
<3 pe:
a =i
es ro)
J Zz
SNI
rc z
= ro)
ye i
2 3 2
2 B= E
O 7
m ;
wn ey =

ES SMITHSONIAN

(op)

= <

fem. a
ernee 5
SN 6
y 2 =

2 c
SNINVINOSHLINS

Ss “0

on w

= oe

S oc

ra) cas

=z ae |
ES SMITHSONIAN

vat re

2) a
, nen :
t\ > a
ja) Ey se
y i” =

Zz a
SNI

ee

et
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